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THE MAIN AIM OF TAXONOMY OF SALTICIDAE, AND THE DUTY OF TAXONOMISTS, IS TO DISCOVER, DEFINE
AND CLASSIFY >4600 INSUFFICIENTLY KNOWN SPECIES OF THESE SPIDERS |
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Abstract
Index. The guide provides short, pictoriall key to the following groups of genera, linked from here : AEURILLINES , AMYCINES , AMYCOIDA VARIA , ASTIAINES , BELIPPINES , CHRYSILLINES, COCALODINES, COLONINES[=THIODININES], DENDRYPHANTINES , DIOLENINES , EUOPHRYINES , EUPOAINES , EVARCHINES, FOSSILS , HABRONATTINES , HARMOCHIRINES , HELIOPHANINES , HISPONINES , HYLLINES, ICIINES , LAPSIINES , LIGONIPEINES , LYSSOMANINES , MENEMERINES, MYRMARACHNINES , NOTICIINES , PELLENINES, PSEUDICIINES, SIMAETHINES , SITTICINES , SPARTAEINES , SYNAGELINES , THIRATOSCIRTINAE ,
, YAGINUMAELLINES, YLLENINES. (Switch
also to interactive index of all genera).
ABSTRACT. The database of Salticidae is divided now into 1) - this Part I entitled "Introduction to alternative classification of Salticidae" giving quick scan of comparative diagnostic data; 2) comparative Part Ia entitled "Maddison's views on classification of Salticidae"and 3) extensive database called the Part II - "Global Species Database of Salticidae (Araneae)" and available at http://www.peckhamia.com/salticidae/index.html .
This Part I gives:
I. Proposal of main diagnostic characters delimiting and defining groups of genera in a practical purpose to facilitate identification.
II. Grouping by these
characters ALL 4568 species
divided into 636 genera (as counted on the day of writing, these numbers change continuously due to discovery of new taxa, and to merging of some taxa), documented by diagnostic drawings and photos in the whole relevant literature. [ATTENTION: Taxons having no diagnostic drawings are not included here and considered species inquirenda, or dubia, they are listed and explained in Part II of the present database. Current World Spider Catalog http://www.wsc.nmbe.ch/, which includes these names, lists 5790 described species, divided
into 602 nominal genera.] .
III. Permits to check stability and diversity of these characters,
including also cases of their misapplication.
IV. Genera are joined by similarity of characters into informal "groups of genera" (some of these "groups" are joined into "supergroups of genera") facilitating practical identification, these are not intended to replace traditional subfamilies and higher categories, although morphological similarities may suggest relationships.
ATTENTION. Due to incomplete
and often unclear documentation in the literature, the proposals of delimitation
have working character, pending further research and tests. Each entry below consist of short definition, thumbnails of
drawings/photos a few exemplary species, and names of group of genera (simultaneously
a link to comparative plates of the best diagnostic charateres for ALL SPECIES of
the group).
DISCLAIMER - BEWARE OF UNAUTHORIZED OR INVALID COPIES. This monograph, freely accessible on Internet, was developed gradually during years 1995-2016, with subsequent versions updated and replaced at six months intervals. In a result various versions, not updated, are circulated,or stored, parallel on some servers, or DVD. In particular, unauthorized and outdated version displayed at http://salticidae.org/salticid/main.htm represents my views prior to 2007, but was quoted as a source even in some papers published in 2015. Therefore I must stress that the only versions authorized now by myself are those accessible at: http://www.peckhamia.com/salticidae/Subfamilies/ (part I) and http://www.peckhamia.com/salticidae/ (part II). |
Introduction
Why new kind of groups of genera?
The most important step in identification of Salticidae, and other groups of animals, is recognition of a genus.
Genus in taxonomy may be defined (delimited) by any sufficiently distinct character, mutual for all species included, stable (that is its diversity does not exceed accepted limits) and different from all other genera. That definitions fits also suprageneric ranks of taxa.
Species within genus, if correctly placed [misplacements are common] are usually similar enough to recognize them as congeners, if their mutual features are memorized. Memorizing of 636 genera is, however, difficult, because similar features repeat in various unrelated groups in various parts of the world - to recall an example of ant-likeness, appearing in various groups of Salticidae (and in several other families as well). Grouping of genera by their similar properties is therefore procedure facilitating identification. That may be superfluous for the best known faunae of Europe and North America, where identification is assisted by numerous atlases and keys, but is highly desirable for less known faunae.
Problem of division into groups of genera become acute when studies concentrate on incompletely, rather poorly known faunae of the tropics, or other insufficiently known parts of continents and archipelagoes. There are practical questions requiring immediate answer: how to place a species into one of earlier described genera? How to decide that it deserves description as a new genus? Into which group should it be classifies? There seems to be three approaches, attempting to answer these questions.
1. Traditional system popularized by works of Simon 1901-1903, modified by Petrunkevitch 1928, and summarized by Bonnet 1959, based on cheliceral dentition, arrangements of eyes, body proportions, distribution of spines and characteristic groups of setae. These characters are artificial, in a sense that are not related to affinities, are imprecise (eyes II "closer" to eyes I, versus "somewhat more distant"), do not account for diversity within groups. A mess, created by that approach is well illustrated in the subchapter Comparison of HISTORICAL classification of Aelurilleae.
Modern authors try to improve the system by listing some exemplary exceptions and additions to particular subfamilies, but without solving the essential problem. They use names of subfamilies, but there are no definitions of many of them and no stated criteria of classification of particular genera.
2. There are attempts to base subdivision of family Salticidae by molecular data, especially by gene sequencing, developed by Maddison at al. (among other 2003b,
2008,
2012,
2014,
2015). A concise (61 pages!) summary of conclusions drawn from that approach was recently published by Maddison 2015 (Part Ia of this database is rearranged according to his ideas). Valuable for study of evolution, as they seem to be, they are not translated yet into practical classification, there are no correlations with observable, morphological characters. ["... In the molecular
phylogeny ... A high genitalic diversity could occur even in closely related species, if for instance strong sexual selection drives rapid divergence..." (Zhang J., Maddison W.P., 2015: 938 (1): 30) - an opinion acceptable for some scale od diversity, but clearly exaggerated in practical application..]. With inductive interpretation, how can we know whether presented data are sufficiently representative for a group they purport to characterize? Another weakness of that approach is that collective units are created by joining together genera, taken oun from various units, without (temporarily perhaps) defining them, that is without indication which morphological characters are mutual to them, and which separate them from other units of the same rank, for instance by which characters we can join into Astioida such diverse tribi as Myrmarachnini, Neonini and Astiini? The credibility of conclusions is not increased by incidents of apparent misidentification of representative species, and some strange pairing of taxa. The contents of tribi in the Maddison's 2015 system - see Part Ia -
"Maddison's views on classification of Salticidae".
3. Comparison of selected, easily noticeable morphological diagnostic characters, such as male palps together with internal structures of epigyne, checked for representability and stability of ALL species known. That approach become possible now, owing to accumulation of diagnostic morphological drawings during
last 60 years by several researchers, using similar methodology and purporting to define particular species and genera. Presented in the Part II of the Monograph - Global Species Database of Salticidae) these are now displayed in orderly comparative fashion in a this Monograph... Part I - "Introduction to alternative classification" (and compared in parallel Part Ia - "Maddison's views on classification of Salticidae"
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permitting to see a pattern of similarities and their gradual changes, amounting to a new system of identification of Salticidae (see below). Although still a temporary compromise, pending complementary research and further reclassifications, it tries to help classification of Salticidae of poorly known geographical faunae (in fact ALL continents except Europe and North America).
4. Mutual end of the three above systems are genera, their grouping purports to help identification and to suggest their relationships. In Simon system genera are joined into groups of genera, Petrunkevitch 1928 named them "subfamilies", which was followed by all subsequent authors. The definitions of particular subfamilies are insufficient and delimitation of genera is subject of endless variations, due to scanty knowledge. Equivalents of subfamilies in Maddison's system are named "tribi" (singular "tribus"), with taxa of higher ranks named "clades", grouped into six "subfamilies" (in a new sense proposed by Maddison). The most speciose of these "subfamilies" is Salticinae (5379 species grouped into 538 genera). The declared purpose of that division is to show phylogenetic relationships between genera, with lip service paid to their identification. In difference to that, the system proposed by Prószyński concentrates on identification of genera, their groupings being functional equivalent of "subfamilies" and "tribi", they may hint also on their relationships but abstaining from premature speculations.
5. Documentation. Presentation of diagnostic features without adequate documentation is insufficient for scientific purposes, translation of appearance of characters into words is too imprecise and often misleading. Diagnostic drawings should be precise and without simplifications, details should be shown in adequate size. Drawings should be verified by color photographs and/or computer enhanced automontage, these are nowadays so easily accessible (also displaying them without expenses in the Internet) that there are no excuses for publication of descriptions without such documentation.
Wide scope of application of male palps for diagnostic purposes resulted from empirical observations. Palps are conservative, stable structures, characterizing large groups of genera, with small modification of details in related genera and species. These are the only structure which, after quick glance, permits identification of subfamily or group of genera. There are no other structures serving that level of identification so well. Examination of palps involve detaching them from the body and fixing in requested standard positions (the easiest way to do that is too push palp partially into sands on the bottom of Petri dish) and examining covered by alcohol, oblique illumination by a micro-lamp helps to differentiate details of structure). The most convenient way of examination of palps is to document them in their resting state, in ventral and retrolateral position (tibial apophysis up). Recent research of several authors indicate that structure of male palps, especially of embolus and conductor, is more complicated than heretofore assumed, it is therefore advisable to make additional studies aimed at better understanding their details and function.
Types of internal structure of epigyne characterize well groups of species within a genus, their fine details help to identify species. There are some traits of structures characteristic for groups of genera, even subfamilies, but these require further comparative studies. An arachnologists must be warned that evolution of spermathecae and ducts runs parallel in not related genera and this may be in some cases misleading (for instance ducts making double spiral in some Marpissa and Yllenus). Examination of internal structures of epigyne may last a day of work (macerating soft tissue, rinsing, staining in Chlorazol Black E, mounting as temporary slide in transparent medium, finally storage after examination in a microvial) but expenditure of time may be shortened by making a series of preparations at a time. Fine details of internal structures should be documented in their true view, without omissions and diagrammatic simplifications, current publications include numerous false synonyms of species and genera due to careless, simplified presentation of these structures.
External appearance of epigyne may help identify species within a genus, but is unreliable for larger groups of species and cannot substitute for its internal structures.
Natural coloration of alive or fresh specimen, displayed on a photo (also dry specimens, preserved like insects, retains their coloration for hundred of years) are probably one of best characters for identification of local species. But WARNING - they are adaptative and are often developed independently in several unrelated genera within the same, or distant areas, even on different continents (for instance patterns of light reflecting, iridescent scales). These properties were unrecognized, when research was conducted on long preserved, faded specimens).
Other body features like cheliceral dentition, spines, bunches of setae, spots of color setae, etc., were popular among arachnologists as easily observable. These can be useful if compared directly, but described obligatorily in a routine manner, as generally practiced, are often useless. Body shape, proportions and size - may be very well memorized, but are difficult and tedious to describe. They are variable to large extent.
Non visible features (based on specialized techniques, used without explanation how they are received and without passing taxonomic control procedures of checking their diversity within each genus and stability), are useless for diagnostic purposes, which are the domain of taxonomy.
. 6. Practical solution to application of subfamilies in Salticidae. Lack of taxonomic revisions of contents of genera, included by different authors into particular subfamilies, together with insufficient definitions and imprecise diagnostic characters of particular taxa, create difficulties in practical application of the subfamily concept in Salticidae. For practical reasons, this series of Keys uses informal GROUPS OF GENERA designed for facilitating identification of genera. There is no provision for usage of GROUPS OF GENERA in the International Code of Zoological Nomenclature, so they can be used informally, however, they were used by Simon 1901-1903 as equivalent of subfamilies and we can follow that tradition provisionally in the present study. Each GROUP OF GENERA is followed by temporary definition and list of comparable taxa used by classical authors: Simon 1901-1903, Petrunkevitch 1928, and Bonnet 1959. The similarity of genera within each group can presumably indicate their affinities, but establishing of affinities is not main purpose of their delimitation. To indicate differences with formal subfamilies, the names of those groups of genera are written in CAPITAL LETTERS and distingushed by ending -ES, instead of formal -INAE.
.7. Procedures of identification of genera, proposed by the Alternative Classification permit to find names of each of over 600 genera by simple, logical steps, available for use by all arachnologists after overcoming psychical barrier of fears of not so difficult preparations: examining palps and clearing and staining internal structures of epigyne. These may create difficulties for beginners, but after initial learning and some practice permit quick identificationus of all Salticidae. The proposed way of identification follows my own process of recognition of unknown species: I usually begin from vague recollection that "have seen something like this already", followed by realization where I could possibly seen it, and then searching for drawings of similar structures in literature. That process lasted weeks in beginning of my career, but searching for in an electronic database shortened it to minutes, and with plates in the present "Introduction to alternative classification ..." is shortened to seconds. Competitive traditional methods of following many details of external body appearances in the Keys following Simon' procedure are very complicated and may be misleading.
In my current practice, the fastest way to check position and relationships of a species is by clicking at the possible genus names in the INDEX of Genera in this "Monograph...", displaying within seconds characters of a set of related species (links marked M show that position also in the Maddison 2015 system). Searching for a an enigmatic species, I follow pictorial Key (below) arranged by palps in their resting status, often looking also on internal structure of epigyne. Since some species differs considerably from their supposed type species (often misplaced in their nominal genera) that searching requires often checking of a number of species.
8. Practical difficulties in applying Alternative Classification
First difficulty is incomplete material, especially from the less studied areas of the world. Presumably large part of Salticidae species is not even discovered, many existing collections are still not described. Position of a species is defined by characters of BOTH males and females, primarily on male palps and internal structure of epigyne, so classification of unmatched specimens is difficult, but there are genera in which no single female is documented.
Diagnostic drawings and photographs, if exist at all, are of various quality. Species of Eastern Hemisphere, studied for 50 years by several taxonomists with comparable methods, are relatively much better known. Photography of alive Salticidae is developing particularly well in Australia, and there is a significant progress in SE Asia and India. Serious harm for knowledge of faunae is inflicted by policy of several Publishers, refusing permission to copy diagnostic drawings and photographs to which they hold copyrights. How the diagnostic features of newly published Salticidae could be digested and included into the classificatory system, if they cannot be compared with series of those already known?
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Definitions of proposed groups of genera of Salticidae
based on palps and epigyne
ATTENTION: taxa mentioned below are defined by diagnostic drawings of ALL species (to see them please follow links).
Proposed division does not include characters based on invisible, non-morphological characters.
Groups of genera standing apart
MENEMERINES
Type genus Menemerus Simon, 1868 , of which type species is M. semilimbatus [Attus semilimbatus Hahn 1827 [1829]) .
Definition. Male palps with robust, fleshy base of embolus, with small, sclerotized embolus atop, often double, in some cases there is soft looking attachment, hidden behind embolus (like in Menemerus bivittatus), of unknown function. Epigyne well sclerotized, ducts in Menemerus and Leptorchestes straight and thick walled, spermathecae located posteriorly. In Kima, however, ducts, are thin and entangled. Menemerus has body relatively flattened and broad, two remaining genera are ant-like, but their carapace is not constricted.
To check diversity of diagnostic characters in ALL species of this group of genera click here!
SYNAGELINES
Interactive
index of genera.Type and only genus Synageles, of which type species is Synageles venator..
Definition. Genus Synageles stands apart from other groups of genera, differing from EUOPHRYINES: in males by lack of meandering spermophor, relation of embolus to inflatable haematodocha is unknown (temporarily?). In females epigyne and its internal structures do not resemble EUOPHRYINES.
To check diversity of diagnostic characters in ALL species of this group of genera click here!
SPARTAEINES
Type genus Spartaeus Thorell, 1891, of which type species is Spartaeus spinimanus [= Boethus spinimanus Thorell 1878]..
Cyrba algerina |
Definition. Considered primitive"Basal Salticidae", are characterized by proportionally enormous, globular or oval spermathecae, in males embolus is robust, arising antero-laterally and encircling tightly anterior part of bulbus. To check diversity of diagnostic characters in ALL species of this group of genera click here!
Supergroup of genera CHRYSILLOIDA
with embolus atop narrowing base, overlaying bulbus,
with only anterior part of spermophor visible
Type group of genera CHRYSILLINES.
CHRYSILLINES
Type genus Chrysilla Thorell, 1887, of which type species is Chrysilla lauta Thorell, 1887.
Phintella versicolor [= Chrysilla v.] |
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Definition. Basis of embolus triangular, overlapping bulbus as a thin, apically narrowing layer, ending by short, thin embolus in front of bulus, laterally to it. Only anterolateral part of bulbus, with part of spermophor loop extends from under that layer. That combination of characters: obliquely running basis of embolus and anterolateral angle of bulbus, emerging from under it, seems to be mark of Chrisillinae. Diversity in exact direction of edge of basis (which may be also tranverse accross bulbus), seemingly lateral position of basis in relation to bulbus, length of embolus and its' bending laterally in a characteristic way, creates difficulty in placement of some genera, and led to separation of groups of genera Iciinae, Noticiinae and Thiodininae, requiring further, more precise studies and, possibly, searching for supporting correlations.
Spermathecae are usually oval or globular, ducts usually straight, running anteriorly, or their derived states.
REMARKS. Chrysillinae is large group of genera, distributed worldwide, it contains several well known, prolific genera considered as types of traditional subfamilies of their own, like Freya, Plexippus, Salticus and other. Acceptation of relationships of oblique basis of embolus to spermophor, emerging from beneath basis of bulbus, leave no other choice of interpretation, unless more convincing morphological correlations would be found. To check diversity of diagnostic characters in ALL species of this group of genera click here!
ICIINES
Type genus Icius Simon, 1876 of which type species is Icius hamatus [= Icelus notabilis Koch C.L., 1846 - preoccupied]
Icius hamatus |
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Definition. Temporarily delimited group of genera, with characters close to Chrysillinae, from which seems to differ by indistinct separate basis of embolus, spermatheca more complicated. Palps in larger genera, like Salticus rather diversified, with habitus more uniform. To check diversity of diagnostic characters in ALL species of this group of genera click here!
COLONINES (FORMERLY "THIODININAE")
Newly designated representative species- Colonus sylvana [= Thiodina sylvana (Hentz, 1846)].
[former type species Thiodina nicoleti Roewer, 1951 has entirely different palpal organ, incompatible with Colonus and its relatives, and should be reclassified to AMYCINES!].
Colonus sylvnus ........................................................................................................................................................Gedea flavogularis |
Definition. Anterior half of bulbus with prominent semiarch of spermophor, posterior half of bulbus fleshy, the border betwen these part runs transversally (sometimes obliquely) across the whole breadth of bulbus. Tibial apophysis usually biramous. Mainly Western Hemisphaera genera. To check diversity of diagnostic characters in ALL species of this group of genera click here!
NOTICIINES
Type genus Holoplatys, of which type species is
Holoplatys planissima [= Marptusa planissima Koch L. 1879].
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Habrocestoides bengalensis |
Definition.
Fleshy
basis of embolus, is developed anterolaterally, apically inclined or bent transversally, pressed
to the anterior edge of bulbus, embolus is characteristically bent
anterior-wards at about mid-line of bulbus. Superficial layer (soft and
fleshy? sclerotized?) developed on ventral surface of bulbus runs diagonally
towards basis of embolus, leaving part of loop of spermophor visible. Mainly
Australian Region , a few genera S & E Asian. To check diversity of diagnostic characters in ALL species of this group of genera click here!
SIMAETHINES
Type genus Simaetha Thorell, 1881, of which type species is Simaetha thoracica Thorell, 1881 .
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Ligurra latidens + Stertinius onoi |
Definition. .Carapace flat and broad, often trapezium shaped, body parts covered with scattered light reflecting scales, or white setae. eye field usually flattened and trapezium shaped, posterior slope of carapace step. Bulbus
simple with spermophor following its contour, basis of embolus intersecting bulbus obliquely, embolus short, arising antero-laterally, short tibial apophysis, Epigyne with prominent median anterior pocket, resembling Harmochirinae, from which differ by spermathecae consisting of two globular chambers separated by connector, ducts short with walls medium thick, running posteriorly. ATTENTION: body shape caused mistakes with Rhene, from which differs by palps and epigyne. To check diversity of diagnostic characters in ALL species of this group of genera click here!
To check diversity of diagnostic characters in ALL species of this group of genera click here!
HELIOPHANINES
Type subgenus Heliophanus Koch C.L., 1833 (Heliophanus) Wesolowska, 1986, of which type species is Heliophanus (Heliophanus) cupreus [= Aranea cuprea) Walckenaer 1802].
Heliophanus curvidens + Heliophanus dampfi |
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Definition. Bulbus has, more or less, triangular outline and is covered by thick and opaque tegument, hidding spermophor, continuouson on embolus, which is therefore immovable - "fixed" to bulbus. Many species have developed lobes or protuberances on bulbus ("bumps" in definition by Maddison). Nominal subgenus Heliophanus (Heliophanus) has prominent process on tibial femur, single or split apically, and two slender apophyses on tibia, other subgenera (Helafricanus) may have apophysis on tibial patella. Females have strongly sclerotized epigyne, plate like or concave, with large central groove, ducts are simple and short, but very strongly sclerotized, spermathecae differ little from ducts. External appearance of males and females is uniform, usually dark, often with some colorless scales scattered, or grouped into white abdominal spots, or semilunar whitish line on anterior edge of abdomen. Small spiders living on vegetation and ground.
REMARKS. Memorized picture of Heliophaninae is bazed on nominal subgenus Heliophanus (Heliophanus), containing 81 species and distributed mainly in Palaearctic Region, remaining less prolific and little known genera, are distributed in Asia and Africa. Numerous genera placed by Petrukevitch 1928 into his compilatory subfamily Heliophaninae do not conform to the definition and are reclassified here to CHRYSILLINES and elsewhere.To check diversity of diagnostic characters in ALL species of this group of genera click here!
Supergroup of genera HYLLOIDA
Groups of genera with embolus arising directly from bulbus, beneath tegulum edge, spermophor translucent, runs marginally, without making any loop in the centre of bulbus
Type group of genera HYLLINES.
HYLLINES
Type genus Hyllus Koch C.L., 1846, of which type species is Hyllus giganteus Koch C.L., 1846.
Hyllus semicupreus |
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................................................................................................... ................................................................................................................... Lystrocteissa myrmex |
Definition. Embolus arises directly from bulbus, usually beneath edge of tegulum. In typical case, point of arising is located near posterior part of bulbus rim, in other embolus arises somewhere on lateral side of bulbus, exceptionally anterolaterally. Embolus makes initial bend at narrow space to bulbus, next running parallel to side of bulbus, close to it, after reaching anterior end of bulbus it continues to run ahead, slightly bent. Embolus is narrow, but not hair thin, in posterior part has narrowing fleshy component. In some cases embolus encircles bulbus entirely. Tibial apophysis usually short, often truncated blunt.
To check diversity of diagnostic characters in ALL species of this group of genera click here!
PSEUDICIINES
Type genus Pseudicius Simon, 1885, of which type species is Pseudicius encarpatus [= Aranea encarpata Walckenaer, 1802] .
Afraflacilla asorotica |
Definition. Embolus of Hylloidea type, this group of genera can be recognized by such somatic characters as subocular row of stridulatory spines, characteristic tibial modification, much longer and robust legs I, body flattened and elongated, color pattern. Epigyne usually with well developed pair of sclerotized pockets, duct usually in form of complicated coils, with prominent armature of scent exuding opening, often developed atop specialized branch of a duct. To check diversity of diagnostic characters in ALL species of this group of genera click here!
PELLENINES
Type genus Pellenes Simon, 1876, of which type species is Pellenes tripunctatus [= Aranea tripunctata Walckenaer, 1802].
Pellenes nigrociliatus + .................................................................................................................................................... Pellenes lapponicus |
Pellenes lapponicus |
Definition. Epigyne
with prominent median pocket separating a pair of sclerotized grooves. Spermathecae and ducts heavily sclerotized and very complicated. Embolus hidden inside protective cover
(see SEM of - Pellenes
lapponicus and tip of embolus of P. nigrociliatus).To check diversity of diagnostic characters in ALL species of this group of genera click here! REMARK. In traditional system subfamily contained also genus Habronattus, here considered separate group HABRONATTINES!
HABRONATTINES
Type genus Habronattus Pickard-Cambridge F., 1901, of which type species is Habronattus mexicanus (syn. Habrocestum m.) (Peckham, Peckham, 1896).
species intermediate - epigyne like Habronattus - palp like Pellenes |
Definition. Embolus is NAKED, but accompanied by prominent, parallel,and long sclerotized tegular apophysis. Epigyne
with narrow aterior pocket followed by prominent median groove, running up to posterior end of epigyne. Spermathecae and ducts heavily sclerotized and very complicated., form a pair
of double spiral strucures (very insufficiently known). The group contains only single, but enormously prolific N American genus Habronattus (over 100 species). Intermediate species Pellenes
lapponicus and P. ostrisnus have ducts resembling Habronattus,
but palps like Pellenes. To check diversity of diagnostic characters in ALL species of the genus click here!
HARMOCHIRINES
Type genus Harmochirius Simon, 1885, of which type species is Harmochirus brachiatus [= Ballus brachiatus Thorell, 1877)].
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Bianor albobimaculatus |
Definition. Palp confronting HYLLOIDA, with embolus hair thin, encircling round bulbus (often truncated anteriorly). Anterior half of epigyne occupied by large, white,
membranous
"window" , bisected incompletely by prominent median pocket. Ducts membranous and usually short, in some genera join spermathecae by characteristic"C" connection. Spermathecae duct-like and entangled, in some genera form compact body, internally convoluted.
External appearance pretty uniform. To check diversity of diagnostic characters in ALL species of this group of genera click here!
EVARCHINES
Type genus Evarcha Simon, 1902, of which type species is Evarcha falcata [= Araneus falcatus Clerck, 1757[1768]].
+Evarcha kochi ............................................................................................................................ . Evarcha patagiata |
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Definition. Differentiated group of genera, recognizable mainly by memorized external appearance, with some species confronting with
general
type of HYLLOIDA, other departing in shape and location of embolus and shape of bulbus. Epigyne with membranous "window", through which
transluce sclerotized spermathecae of different shape; there is always a pair of pockets posteriorly, visible externally or hidden beneath sclerotized
tegument.
REMARK. Diversity of species suggests advisability of transfers between genera, or even splitting and redefining of this group of genera.
To check diversity of diagnostic characters in ALL species of this group of genera click here!
YAGINUMAELLINES
Type genus Yaginumaella Prószynski, 1976 , of which type species is Yaginumaella striatipes Simon, 1868.
Yaginumaella striatipes |
Yaginumaella senchalensis |
DIAGNOSIS. Characterized by presence of a pair of external pockets on surface of epigyne, in majority of species small, varying in position, in a few species making broad, collar lobes on sides of copulatory openings. Copulatory openings in form of anterior, diagonal slits. Copulatory ducts broad, run almost straight posteriorly, near end of epigyne pass into a few narrower loops of spermathecae. Palpal organs with bulbus oval or narrowing posteriorly, in some (Y. stemmleri) with anterior protuberance. Embolus vary in length, in majority of species naked, parallel to bulbus, in type species inside boader sheath [?]. To check diversity of diagnostic characters in ALL species of this group of genera click here!
YLLENINES
Type genus Yllenus Simon, 1868, of which type species is Yllenus arenarius Simon, 1868.
Definition. Recognizable by possession of adaptative "scopula" brush of ventral setae on tarsus I, serving for quick submerging spider in sand, also shape of carapace with long slope beginning immediately behind eyes III. Diagnostic characters of palps and epigyne display exuberant evolution, with embolus and bulbus confronting to HYLLOIDA in some species, in other unique by development of enormous "conductor", strangely shaped tibial apophysis and crazy looking cymbium. In epigyne ducts evolving from simple, strongly sclerotized, through thin walled loops, to unique, elegant double spiral. To check diversity of diagnostic characters in ALL species of this group of genera click here!
Supergroup of genera AMYCOIDA
Western Hemisphere jumping spiders, represented in Palaearctic Region
by single group of genera SITTICINES. Resembling HYLLOIDA by naked embolus arising from bulbus, but spermophor makes central loop, tooth on anterolateral edge of chelicerae multicusp.
Type group of genera AMYCINES.
SITTICINES
Type genus Sitticus Simon, 1901, of which type species is Sitticus terebratus [= Attus terebratus (Clerck, 1757 [1758]).
Chelieral dentition in Sitticus terebratus + palps and internal structures of epigyne in Sitticus floricola. |
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Definition. Recognizable by prominent "S" shaped central loop of spermophor, continuing later marginally before entering embolus. Embolus thin, encircling bulbus partially or entirely. Another striking character is saw-like multicusps tooth on anterior median edge of chelicerae, while posterior edge and its tooth are not developed (an apparent modification of the cheliceral teeth state in other AMYCOIDA). Spermatheca developed from oval to elongate bent structure, thick walled, joined in the middle with weakly sclerotized ducts, either simple and bent, or forming coils of various degree of complication.
REMARKS. Taxonomic characters of SITTICINES (as defined in present usage) agree with conclusions from gene sequencing research, and geographic evolution of the group, in which only genus Sitticus underwent intensive species radiation in Palaearctics, with a few species returning ultimately to Norh America. To check diversity of diagnostic characters in ALL species of this group of genera click here!
AMYCINES
Type genus Amycus Koch C.L., 1846 of which type species (dubius) is Amycus igneus (Perty, 1833) [= Salticus igneus Perty 1833:].
Reference species - Amycus pertyi Simon, 1900].
Definition. Diversified, poorly known Western Hemisphaere genera. Palps in some have "S" shaped loop of spermophor and SITTICINES like cheliceral
dentition, but
these characters are variable and may depart from the Sitticinae model. Body shape variable, some genera are ant-like. In females copulatory ducts often twisted
around spermathecae, in some genera as double spiral. To check diversity of diagnostic characters in ALL species of this group of genera click here!
AMYCOIDA VARIA - group of genera
Type genus - not designated
Definition. Temporary
grouping of poorly known Western
Hemisphere genera, related to AMYCINES by molecular research
(Maddison and al., ), pending taxonomic generalization. To check diversity of diagnostic characters in ALL species of this group of genera click here!
Supergroup of genera EUOPHRYOIDA
Subfamilies with "free" embolus sitting atop membranous anterior haematodocha (Maddison 1996), structure proven
in some genera, suspected in others, seems to be one of major taxonomic divisions in Salticidae.
EUOPHRYINES
Type genus Euophrys Koch C.L., 1834 of which type species is Euophrys frontalis [= Aranea frontalis Walckenaer, 1802 [Platnick: "preoccupied by Olivier, 1789, but amply protected by usage"].
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Definition. Genera characterized by both: in males embolus twisted into single coil or multi coil spiral, in some cases reduced to incomplete bend, in females by a pair of usually simple, globular
spermathecae. Position of coil of embolus variable, usually in either side, or ventral surface of anterior bulbus, in some cases, however, on posterior part of lateral surface (Attention: latest photographic documentation of some genera show double "embolus" structure, which requires further consideration). Associated character is meandering course of spermophor, consisting of two, or one and half incomplete bends, on retrolateral surface of bulbus, usually extending onto ventral surface. Epigyne with two white, membranous "windows", sometimes fused into single one, with translucent globular spermathecae, ducts often simple, running anteriorly or laterally. ATTENTION: Newly proposed group EUODENINES houses genera with internal structures of epigyne resembling DENDRYPHANTINES (copulatory openings located anteriorly, initial part of ducts running posteriorly and then passing into a number of coils, a knot, or a compact body with internal convoluted ducts and with palps having straight or only slightly waving. Also genera with large coil of embolus located laterally on bulbus. Of the other hand genera with palps and epigyne resembling Ballus are placed in group BALLINES, (following Benjamin, 2004). To check diversity of diagnostic characters in ALL species of this group of genera click here!
BALLINES
Type genus Ballus Koch C.L., 1850[1851] , of which type species is Ballus chalybeius (Walckenaer, 1802).
EUOPHRYINES like genera, characterized by epigyne with anterior copulatory openings, ducts straight running along most of lengt of epigyne, with spermathecae in form of enangled coils posteriorly. Palps with embolus coiled atop bulbus, and spermophor stretching almost straight along retrolateral surface of bulbus. Legs I with femur and tibia broadened and dark, metatarsus and tarsus narrower and light. Carapace dorsally flattened..To check diversity of diagnostic characters in ALL species of this group of genera click here!
DENDRYPHANTINES
Type genus Dendryphantes Koch C.L., 1837, of which type species is Dendryphantes hastatus [= Araneus hastatus Clerck 1757 [1758].
Definition. Genera with embolus atop membranous anterior haematodocha, in front of
bulbus; embolus short, straight or branched, single or split, but usually not coiled. Spermophor after initial anterolateral bend runs almost straight along lateral surface of bulbus. Epigyne heavily
sclerotized, with a pair of lateral, or anterior depressions, with copulatory openings at the bottom, often
connected by semi-lunar furrow, copulatory ducts broad, running posteriorly,
spermathecae in a form of entangled loops or compact body internally convoluted.
To check diversity of diagnostic characters in ALL species of this group of genera click here!
EUODENINES Genera with palps and epigyne intermediate between EUOPHRYINES and DENDRYPHANTINES
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The group contains genera intermediate between EUOPHRYINES and DENDRYPHANTINES [hence name EUOPHRYINE-DENDRYPHANTINE] and consists of species resembling DENDRYPHANTINES - with ducts running along the whole length of epigyne, from anterior openings to posterior coils of spermathecae and with coil of embolus atop membranous haematodocha and spermophor gently waving or straight. Details of these structures are often insufficiently documented, unclear or, presumably highly evolved, departing from simple general diagram. Group pending further considerations.To check diversity of diagnostic characters in ALL species of this group of genera click here!
AELURILLINES
Type genus Aelurillus Simon, 1884, of which type species is Aelurillus v-insignitus [= Araneus litera v insignitus Clerck 1757 [1758]].
Definition. Bulbus covered entirely by thick, opaque tegulum through which spermophor does not transluce, embolus coiled behind anterior part of tegulum, with only tip emerging. Tibial apophysis usually split into two rami, usually short, in Langona tibia is single but accompanied ventrally by a bunch of stiff setae. Spermathecae usually compact body convoluted inside, or consisting of complicated chain of chambers (like in Phlegra or Langona). Robust,
terrestrial forms, running or ambushing. To check diversity of diagnostic characters in ALL species of this group of genera click here!
MYRMARACHNINES
Type genus Myrmarachne MacLeay, 1839, of which type species is Myrmarachne melanocephala MacLeay, 1839.
Myrmarachne tristis |
Definition. Genera recognizable by parallel "pipes" like spermathecae with membranous large but irregular ducts attached to their posterior end [the latter visible ONLY on cleared slides, stained with Chlorazol Black E]. Males with palps uniform, bulbus encircled twice by embolus, usually with thin, incomplete "additional" loop of spermophor. Tibial apophysis twisted cork-driver like in Myrmarachne, in other genera also short but straight. In males chelicerae enormously elongated, with multiple isolated teeth (true "pluridentati"). Body usually ant-like, with carapace also constricted; since body appearance is adapted to mimicking ants, same habitus may repeat in independent species. To check diversity of diagnostic characters in ALL species of this group of genera click here!
LIGONIPEINES
Type genus Ligonipes Karsch, 1878
, of which type species is Ligonipes illustris Karsch, 1878.
Definition. Tibia Ist broadened and flattened, almost circular, dark with dense setae and robust spines. Spermathecae with central part "pipe" like, as in MYRMARACHNINES, Male palps intermediate between MYRMARACHNINES and EUOPHRYINES, bulbus with modified meandering spermophor, coil of embolus modified, broadened and encircling bulbus. Male chelicerae not expanded, cheliceral dentition - multicusp saw-like tooth on swollen retrolateral edge of chelicerae.. To check diversity of diagnostic characters in ALL species of this group of genera click here!
BELIPPINES
Type genus Belippo Simon, 1909 of which type species is Belippo anguina Simon, 1909.
Definition. BELIPPINES have exceptional tibial apophysis in a form of sclerotized, short sickle, sitting atop transparent membranous basis - characterized in descriptions as "movable", its function remain a mystery. Bulbus round, encircled with hair thin embolus, spermophor along contour of bulbus with small additional, flattened loop. Females differing from other ant-like subfamilies by epigyne with pipe-like spermathecae, expanded anteriorly into globular vesicle, copulatory ducts membranous and visible only in cleared and stained preparations, in forms of very complicated loops. Abdomen and carapace not constricted, pedicel short. To check diversity of diagnostic characters in ALL species of this group of genera click here!
HISPONINES
group of genera of uncertain
status
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................................................................................................... .............................................................................. Lystrocteissa myrmex - see HYLLINES |
- note resemblance to fossil Prolinus by location of eyes anterior lateral and posterior median
(the latter strikingly reduced in size)
on
the same protuberance in extant Tomocyrba, Hispo,
some "Massagris" and Lystrocteissa - interpreted as relict. To check diversity of diagnostic characters in ALL species of this group of genera click here!
DIOLENINES
Type genus Diolenius Thorell, 1870 of which type species is Diolenius phrynoides [= Attus phrynoides Walckenaer, 1837].
Definition. Narrower specialized group of genera, with distinctly (sometimes enormously) elongated trochanter and coxa I. Palpal organ derived (according to gene sequencing data) but not obviously similar to Euophryinae, tip of embolus emerging anteriorly from behind bulbus. To check diversity of diagnostic characters in ALL species of DIOLENINAE click here!
Groups of genera with simple palps and
epigyne
COCALODINES
Type genus Cocalodes
Pocock, 1897, of which type species is Cocalodes leptopus Pocock, 1897.
Definition. Bulbus with simple sclerotized apophysis, spermophor following contour of bulbus, embolus short, arising antero-laterally. Epigyne with simple spermathecae,
ducts short, thick walled .Poorly
known and ill defined, live in South Eastern Hemisphere. To check diversity of diagnostic characters in ALL species of this group of genera click here!
LAPSIINES
Type genus Lapsias Simon, 1900, of which type species is Lapsias estebanensis Simon, 1900.
Definition. Bulbus
with simple sclerotized apophysis, spermophor following contour
of bulbus, embolus encircling bulbus entirely or parially. Epigyne with complicated
spermathecae and ducts. Poorly
known and ill defined, live in South Western Hemisphere. To check diversity of diagnostic characters in ALL species of this group of genera click here!
ASTIAINES
Type genus Astia Koch L., 1879
, of which type species is Astia hariola Koch L., 1879
.
Definition.
Bulbus simple oval, embolus short, arising
antero-laterally. Epigyne
with small, simple spermathecae, ducts thin, running anteriorly, cheliceral inner
posterior tooth - saw-like (numerous small cones arising from mutual basis).Poorly known and ill defined, live in South Eastern Hemisphere. To check diversity of diagnostic characters in ALL species of this group of genera click here!
Pending placement
EUPOAINES
Type genus Eupoa Żabka, 1985 of which type species is Eupoa prima Żabka, 1985.
Eupoa nezha + Eupoa prima + Eupoa sp from Thailand |
Tarne dives |
Definition. According to Maddison 2006 presence
of a median apophysis in the male palps and of a tarsal claw in the female palps place Eupoa inside Basal Salticidae. My own knowledge is insufficient to comment upon that yet. Pending further studies. To check diversity of diagnostic characters in ALL species of this group of genera click here!
Taxon of uncertain position - Thiratoscirtinae
a new clade of subfamily rank, proposed by Bodner & Maddison 2012, p. 221-222),
with taxonomic documentation limited to a list of genera included.
(Use Index to find data of proposed genera).
Infrafamily LYSSOMANINES
Originally defined by eyes arranged in four rows, however, there are unrelated genera with similar eyes
arrangement, or intermediate to three rows, palps and epigyne are insufficiently studied
LYSSOMANINES
Type genus Lyssomanes Hentz, 1845, of which type species is Lyssomanes viridis [= Attus viridis Walckenaer, 1837].
b)c)
Asemonea tenuipes + b) Lyssomanes viridis + c) Lyssomanes longipes |
b)c)
Asemonea tenuipes + b) Lyssomanes viridis + c) Lyssomanes longipes |
Definition. Eyes arranged in 4 rows, the
2nd (anterior lateral) distinctly above the 1st (anterior median). Prolific genus Lyssomanes is known from the Western Hemisphere (allmost all species in Central and South America). In S and S Asia only Asemonea and Hindumanes Several genera have eyes arranged in four rows (or intermediate between four and three rows( but their palps and epigyne are incompatible so should be reclassified elsewhere. These are Athamas, Leptathamas (Euophryinae), Onomastus, Orthrus (Astieae). To check diversity of diagnostic characters in ALL species of this group of genera click here!
ADDITION - MATERIAL FOR THINKING
FOSSILS
b)
Eolinus tystschenkoi +b) Baltic sp [photo Hill] Baltic amber fossils. |
Fauna
of the past - amber preserved Eocene Salticidae - these
specimens are remnants of Tertiary pine forests fauna, preserved in amber resins
leaking from trees, found, for example, in sands submerged in the Baltic Sea
and inland. REMARKS. Affinities with tropical extant fauna assumed, but not precisely identified. To check diversity of diagnostic characters in ALL species click here!
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