Jonathan C. Harrod, Herbert W. Levi, and Laura B. Leibensperger.
The neotropical orbweavers of the genus Larinia (Araneae: Araneidae).
Psyche 97:241-266, 1990.

Full text (searchable PDF, 3336K)
Durable link: http://psyche.entclub.org/97/97-241.html

The following unprocessed text is extracted from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

THE NEOTROPICAL ORBWEAVERS OF THE GENUS
LARINIA (ARANEAE: ARANEIDAE)*
BY JONATHAN C. HARROD, HERBERT W. LEVI AND LAURA B. LEIBENSPERGER
Harvard College and Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02 138 This paper is one in a series of publications on Neotropical orb- weavers (Levi 1968, 1971, 1985, 1986, 1988, 1989a, 1989b, in press), made to facilitate identification of common American spiders. It is hoped that this information will also contribute to our knowledge of the phylogeny and relationships of orb-weaving spiders. Larinia consists of 30 to 40 species, the genus having world-wide distribution. In the 1970s, Grasshoff (1970a, b, c, 1971) revised the African and Indoaustralian species of Larinia, placing them into nine genera. Genera were distinguished by details of palpal struc- ture, especially of the radix, (a projection of the tegulum, illustrated here in Fig. 41). Grasshoff (1971) placed the American species Lari- nia directa in the genus Drexelia, this genus being distinguished by the internal structure of the epigynum: ducts that are wide on the outside and narrow toward the seminal receptacles (Levi 1975, fig. 9, 12). Levy (1986) also used Grasshoffs genera in revising Larinia of Israel. Alternatively, Levi (1975), revising the Larinia of North America, placed all species in Larinia, as did Marusik (1986), with Larinia of the USSR and Tanikawa (1989), with Larinia of Japan. In the current paper, we use Larinia in this broad sense, including the nine genera into which Grasshoff split the genus. We do not follow Grasshoff, partly because of the difficulty in placing our species known only from one sex into his genera. Operationally, we consider Grasshoffs genera to be species groups. Larinia has several synapomorphic characters, including an elon- gate abdomen, (in a family whose species usually have a spherical abdomen), the anterior median tubercle-like point of the abdomen, a white median ventral streak (also found in the unrelated araneid genera Eustala and Metepeira), and the structure of the epigyna and palpi (see below).
*Manuscript received by the editor August 24, 1990. 24 1
Pu&e 97:241-265 t HIM), http:llpsyche cnlclub orBf97Μφ7-24 hlml



================================================================================

242 Psyche [vo~. 97
Levi (1975) revised and illustrated the three North American spe- cies of Larinia: L. directa (Hentz), L. borealis Banks and L. famula- toria (Keyserling). All three are found in Mexico; L. directa is also found throughout the Neotropics. This paper includes the nine spe- cies found in the West Indies and south of Mexico. Materials and Methods. Methods are similar to those described in previous papers. Levi (1977) includes a description of the working method, which will also be discussed in a forthcoming revision of the genus Wixia.
Eye sizes were described using the diameter of the anterior median eyes as a standard. For the anterior eye row, distance between eyes was measured in number of anterior median eye diameters; for the posterior eye row, in number of posterior median eye diameters.
We thank the curators of the following collections for the loan of specimens:
AMNH
BMNH
CAS
IRSNB
MACN
MAES
MBUCV
MCN
MCZ
MEG
MIUP
MLP
MNHN
American Museum of Natural History, New York; N. Platnick, L. Sorkin
British Museum (Natural History), London, England; P. Hill yard
California Academy of Sciences, San Francisco; W. J. Pulawski, D. Ubick
Institut Royal des Sciences Naturelles de Belgique, Brus- sels, Belgium; L. Baert
Museo Argentine de Ciencias Naturales, Buenos Aires, Argentina; E. A. Maury
J. M. Maes, Leon, Nicaragua
Museo de Biologia, Universidad Central, Caracas, Vene- zuela; J. Racenis
Museu de Cigncias Naturals, Porto Alegre, Brazil; A. Lise, E. Buckup
Museum of Comparative Zoology, Cambridge, Mass. M. E. Galiano, Buenos Aires, Argentina
Museo de Invertebrados, Universidad de Panama, Pan- ama; D. Quintero A.
Museo de Universidad National, La Plata, Argentina; R. F. Arrozpide
Museum National d'Histoire Naturelle, Paris, France; J. Heurtault




================================================================================

19901 Harrod et a1.-Neotropical Larinia 243 MHNSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; D. Silva D.
MNRJ
Museu Nacional, Rio de Janeiro, Brazil; A. Timotheo da Costa
MZSP Museu de Zoologia da Universidade de Siio Paulo, Brazil; P. Vanzolini, L. Neme, J. L. M. Leme NRMS Naturhistoriska Riksmuseet, Stockholm, Sweden; T. Kronestedt
SMF Forschungsinstitut Senckenberg, Frankfurt, West Ger- many; M. Grasshoff
USNM National Museum of Natural History, Smithsonian Institution, Washington, D.C.; J. Coddington ZMK Zoologisk Museum, K$benhavn, Denmark; H. Enghoff Research was started with the support of National Science Foun- dation grant GB-36161.
We also thank E. Buckup, who identified a number of specimens in the MCN collection for us, using our preliminary drawings. P. Sierwald read the manuscript and made numerous valuable sugges- tions. D. Silva D. was helpful in locating a Peruvian place name. Larinia Simon
Larinia Simon, 1874: 1 15. Type species by monotypy Epeira lineata Lucas, 1846. The gender of the generic name is feminine. Levi, 1975: 102. Drexelia McCook, 1892: 127. Type species by monotypy Epeira directa Hentz (1847). Diagnosis.
Larinia differs from the related genus Araneus by the elongate, oval abdomen, often with an anterior, median tubercle (Figs. 3, 8, 12, 18, 28, 38,45) and sometimes projecting behind and above the spinnerets (Figs. 19, 23, 29). The abdomen usually has a dorsal pattern of longitudinal marks (Figs. 3, 8, 12, 18, 22, 28, 34, 38,45), and venter with median white markings on black (Figs. 4,9, 13, 19, 23, 29, 35,39).
Eustala fuscovittata and some other Eustala resemble Larinia in shape and markings. However, the genitalia differ: female Eustala have a scape that projects anteriorly from the base, and males have a white colored, lightly sclerotized, cone shaped median apophysis. The resemblance in shape and markings is a homoplasy. Description. The light colored carapace may have a double, black, longitudinal line behind the eyes that fuses into a single median longitudinal line (Figs. 12, 18, 28,45). The sternum may be



================================================================================

244 Psyche [vo~. 97
all black (in L. ambo and L. lampa), all yellow (in L. bivittata, L. directa, and L. t-notata), or yellow in the center and darker around the edges (L. minor, L. montecarlo, L. neblina, and L. tucuman). The legs and dorsum of the abdomen are usually light yellow- orange. The anterior median eyes are the largest, usually separated by a diameter or more. The posterior medians may be slightly smaller, are separated by their diameter at most, and are often in contact. The ocular quadrangle is narrower behind than in front. Males have a tooth on the lateral margin of each endite, lack the hook on the first coxae which is found in some araneid genera, and have two macrosetae on each palpal patella (Levi 1975, Fig. 3). The second tibia is thicker than the first.
Genitalia. The epigynum is lightly sclerotized, and has a wrinkled scape enclosing a pocket, with a lip at its tip (Figs. 1,6, 10). The lip and pocket are synapomorphies shared with Araneus. An exception is L. montecarlo (Fig. 43), but here the scape may have been torn off in all available specimens. The internal female genita- lia were illustrated in a previous paper (Levi 1975). The epigynum frequently contains one or two broken embolus tips from a male palpus, which are difficult to remove (Fig. 27, right side; Figs. 32, 33, on both sides in depressions).
Structure of the palpus is similar to that of Araneus and Nucte- nea. The conductor sits on the rim of the tegulum and there is no paramedian apophysis, (this is usually true of genera with the con- ductor in this position). It is not known whether this condition is primitive or secondary. The presence of two palpal patellar setae is shared with Araneus, Nuctenea and Neoscona. The palpal embolus is heavily sclerotized. In L. directa, it is a diagonal rod (Fig. 5); in L. montecarlo, a horizontal curved rod (Fig. 47). It is triangular in L. tucuman (Fig. 14), anchor-shaped in L. bivittata (Fig. 24), and comma-shaped in L. ambo (Fig. 30). In L. t-notata it is hidden behind a lamella (Figs 40,42). In all species, the tip of the embolus rests on the lightly sclerotized conductor (Figs. 5, 14, 24, 30,47). The conductor is attached to the rim of the tegulum as a white irregularly shaped cushion (as it is in Araneus, Levi in press, fig. 3) in L. tucuman (Fig. 19, L. bivittata (Fig. 25), L. ambo (Fig. 31), and L. montecarlo (Fig. 48).
The most visible structure of the palpus is the median apophysis, lying proximally to the embolus. In L. directa, (Fig. 5), L. tucuman



================================================================================

19901 Harrod et a1.-Neotropical Lurinia 245 (Fig. 14), and L. ambo (Fig. 30), the median apophysis has a distal, sclerotized spine and a basal projection which is blunt and less heavily sclerotized. In L. bivittata (Figs. 24, 25), and L. montecarlo (Figs. 47,48), the median apophysis has two spines. In L. t-notata, the median apophysis is flattened (Fig. 40) with a distal serrated edge (Figs. 41, 42) and a large proximal, pointed, curved prong (Figs. 41, 42). The tegulum of this species also has a flat projection with a serrated edge (right side, Figs. 40,41,42). Distal to the embolus are structures which are difficult to homol- ogize, the terminal and subterminal apophyses. In L. directa, the terminal apophysis is triangular and pointed (Fig. 5). In L. tucu- man, it appears as a "hanging" rod (Fig. 14). In L. bivittata, both the subterminal and terminal apophyses are pointed6'above" the embo- lus (Fig. 24). In L. ambo, they are pointed and almost parallel to the embolus (Fig. 30). In L. neblina, they are two blunt sclerites (Fig. 40, 42). In L. montecarlo, there are two pointed structures, one pointing "up" and the other "down" (Fig. 47). In all cases, there is a hemato- docha between the embolus and the sclerotized portions of the sub- terminal and terminal apophyses. This is small in L. t-notata (Fig. 42).
Distribution. The genus is found world-wide. There are between 30 and 40 species.
Natural History. Specimens are commonly collected by sweep- ing vegetation. Larinia directa sits in the hub of the web at night, on vegetation to the side of the web in daytime (Levi 1975). Drexelia scriba Mello-Leitiio, 1940: 203 from Colatina, Espirito Santo, Brazil, in MNRJ, lost.
Larinia albosigillata Mello-Leitiio, 1947b: 247, is an immature A raneus.
Larinia bristowei Mello-Leitgo, 1940: 180, belongs in a new genus.
Larinia coamensis Petrunkevitch, 1930: 335, figs. 221-224, Q, is Metazygia crewi (Banks). NEW SYNONYMY
Larinia maulliniana Mello-LeitZio, 195 1 : 33 1 is Zygella x-notata.



================================================================================

246 Psyche
[Vol. 97
Synonymized by Levi (1974: 276).
Larinia mundula (Keyserling), 1892: 179 is a Metazygia. Larinia mundulella (Strand), 1916: 1 14 is a Metazygia. Larinia nobilis Mello-Leitgo, 1944: 33 1 is Araneus corporosus (Keyserling). Synonymized by Levi (in press). Larinia rubroguttulata Keyserling, 1880: 3 14 is a Eustala. Larinia silvestris Bryant, 1942: 5 is a Cyclosa. Key to Larinia species, females
1
Epigynum with scape; venter of abdomen with black or gray markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 -
Epigynum without scape, as in Figs. 43,44; venter of abdomen without black or gray markings (Fig. 46); southern Brazil Map 1. Distribution of Larinia directa.




================================================================================

19901 Harrod et a1.-Neotropical Larinia 247 (Map 2). .................................. montecarlo ............... Scape longer than broad (Fig. 32,36). 3
Scape about as broad as long, or broader than long (Figs. 1,6, 10, 16, 20,26). ...................................... 4 Abdomen more than twice as long as wide, with prominent anterior median tubercle; dorsum lightly colored, sometimes with a double row of black spots; epigynum as in Figs. 36,37, with scape often curving to one side; total length greater than ..... 6.0 mm; southern Brazil, Argentina (Map 2). t-notata
Abdomen with length about one and one half width, rounded, without prominent tubercle; dorsum of abdomen with broad brown bands (Fig. 34); epigynum as in Figs. 32, 33; total ....... length usually less than 6.0 mm; Venezuela (Map 2). ........................................... neblina ..
Abdomen more than twice as long as wide (Figs. 3, 18). 5
Abdomen less than twice as long as wide (Figs. 8, 12, 22, 28). ............................................... 6 Distal lip of epigynal scape wide (Fig. 1); total length greater than 6.0 mm; widespread (Map 1). ............... directa Scape with narrow lip (Fig. 16); total length usually less than 6.0 mm; Greater Antilles (Map 2). ................. minor Dorsum of abdomen with a pair of dark lateral stripes, straight or undulating, sometimes with a thin median stripe between them (Figs. 8, 12, 22); epigynum as in Figs. 6, 10, 20. ................................................ 7 Dorsum of abdomen with a single dark toothed median band ........ (Fig. 28); epigynum as in Fig. 26, 27; Peru (Map 2). ............................................... ambo ... Edges of lateral dorsal bands undulating (Figs. 12, 22). 8
Lateral bands with straight edges and filled with red-brown dots; epigynum as in Fig. 6; Peru (Map 2). ......... lampa Total length less than 4.7 mm; dorsal bands light anteriorly, becoming darker towards posterior; pronounced median notch in posterior lip of epigynal base (Fig. 11); Argentina (Map2). .................................... tucuman Total length greater than 5.2 mm; dorsal bands dark through- out entire length. Posterior lip of epigynal base dark, swollen, and sclerotized (Fig. 21); southern Brazil, Chile to Argentina (Map 2). ..................................... bivittata



================================================================================

Psyche
[Vol. 97
Key to Larinia species, males
Palp with both tegulum and median apophysis darkened and with serrated borders, as in Figs. 40-42; total length greater than 5.3 mm; southern Brazil to Argentina (Map 2). ....... ............................................. t-notata Tegulum and median apophysis without serrated borders; .........................
total length less than 5.3 mm.
2
Embolus triangular (Fig. 14), anchor-shaped (Fig. 24), or comma-shaped (Fig. 30). Abdomen less than twice as long as .............................................. wide. 3
Embolus thin and tubular as in Figs. 5, 47. Abdomen more ...........................
than twice as long as wide.
5
Embolus comma-shaped; palpus as in Figs. 30, 31; dorsum of abdomen with a single dark toothed median band; Peru (Map 2). ............................................ ambo Embolus anchor-shaped (Fig. 24) or triangular (Fig. 14); dor- sum of abdomen with a pair of dark lateral stripes separated .........................
by a thin white median band.
4
Embolus anchor-shaped, dark, and sclerotized; terminal apo- physis with two distinct tips (Fig. 24); dark dorsal bands run- ning entire length of abdomen; southern Brazil, Chile to ...........................
Argentina (Map 2). bivittata
Map 2.
Distribution of Larinia species.




================================================================================

19901 Harrod et a1.-Neotropical Larinia 249 -
Embolus triangular, not sclerotized; two tips of terminal apo- physis not distinct; palp as in Figs. 14, 15; dorsal bands light or absent anteriorally, becoming darker towards posterior; Argentina (Map 2). .......................... tucuman 5(2) Median and terminal apophyses as in Fig. 5; tubular embolus lying diagonal to main axis of palp; usually greater than 4.3 mm in total length; widespread (Map 1). .......... directa -
Median and terminal apophyses as in Fig. 47; with tubular embolus bending "down;" usually less than 4.3 mm in length; southern Brazil (Map 2). .................... montecarlo Larinia directa (Hentz)
Figures 1-5; Map 1
Epeira directa Hentz, 1847: 478, pi. 31, fig. 21, Q. Type specimens from South Carolina and Alabama, destroyed.
Epeira tetragnathoides 0.P.-Cambridge, 1889: 16, pi. 7, figs. 9, 10, 9, 3. Male and female syntypes from Guatemala and Panama, in BMNH. Name preoccupied by Epeira tetragnathoides Walckenaer.
Epeira intercisa O.P.-Cambridge, 1889: 18, pi. 5, fig. 11, 3. Male holotype from Bugaba, Panama, in BMNH, lost. First synonymized by F.P.-Cambridge, 1903.
Drexelia directa: - McCook, 1892: 127. F.P.-Cambridge, 1903: 461, pi. 43, figs. 12, 13, $,a. Grasshoff, 1971: 93, fig. 45, Q. Epeira deludens Keyserling, 1893: 261, pi. 13, fig. 195, $2, 3. Syntypes from Florida and Guatemala, in USNM, examined. First synonymized by F.P.-Cambridge, 1903.
Larinia bellona Banks, 1898: 257, pi. 15, fig. 6, Q, 3. Syntypes from Tepic, Mexico, in MCZ, examined. First synonymized by Levi, 1975. Larinia directa: - Comstock, 1913: 508, fig. 545. Levi, 1975: 105, figs. 1-12, 31, 34, 37-41, Q,3, Map 1.
Larinia cymotypa Chamberlin, 1924: 649, fig. 93, 9. Female holotype from Gulf of California, in CAS, examined. First synonymized by Levi, 1975. Larinia albonigra Franganillo, 1934: 158, 3. Male holotype from Cuba, in Cuban Acad. Sci. First synonymized by Levi, 1975. Drexelia octopunctata di Caporiacco, 1955: 349, Q. Female holotype from Caracas, in MBUCV, examined. First synonymized by Levi, 1975. Larinia nigrovittata Mello-Leitiio, 1947a: 12, figs. 26, 27, 3. Male holotype from Carmo do Rio Claro, Minas Gerais, Brazil, in MNRJ, examined. First syn- onymized by Levi, 1975.
Description.
Female from Veracruz, Mexico. Carapace, che- licerae, endites, coxae, sternum, legs yellow. Carapace with orange longitudinal stripe; sternum with gray border. Labium brown. Dor- sum of abdomen with white median stripe, bordered by faint gray



================================================================================

250 Psyche [vo~. 97
bands with four black dots along their lateral margins (Fig. 3); venter with median white stripe bordered laterally by black bands (Fig. 4). Posterior median eyes 0.8 diameters of anterior medians, laterals 0.8 diameters. Anterior median eyes 1.3 diameters apart. Posterior median eyes 0.2 diameters apart. Abdomen oval, more than twice as long as wide, with anterior median tubercle (Fig. 3). Total length 7.4 mm. Carapace 2.7 mm long, 1.5 wide. First femur 3.2 mm, patella and tibia 4.4, metatarsus 4.0, tarsus 1.4. Second patella and tibia 4.0 mm, third 2.1, fourth 3.8. Male from Veracruz. Color as in female. Posterior median eyes 0.7 diameters of anterior medians, anterior laterals 0.5 diameters, posterior laterals 0.6. Anterior median eyes 1.2 diameters apart. Posterior median eyes in contact with each other. Abdomen oval. Total length 5.2 mm. Carapace 1.9 mm long, 1.5 wide. First femur 3.2 mm, patella and tibia 4.7, metatarsus 4.9, tarsus 1.5. Second patella and tibia 4.2 mm, third 1.9, fourth 3.5. Illustration. Specimens from Surinam were illustrated. Variation.
Females varied between 6.2 and 10.5 mm in length. Males varied between 4.2 and 5.2 mm.
Diagnosis.
Larinia directa differs from other species south of Mexico in the wide distal lip on the epigynal scape (Fig. 1) and the tubular embolus lying diagonal to the main axis of the palp (Fig. 5). In Mexico two similar species are found: Larinia famulatoria (Keyserling) and L. borealis Banks. Both are illustrated and keyed out by Levi (1975).
Distribution.
Southern United States through Paraguay. Records. United States localities shown are from Levi (1975, Map 1). MEXICO Baja California: La Ribera (CAS); San Felipe (AMNH); Santa Anita (CAS). Islas Revillasgegidos: Isla Socorro (CAS). Sonora: 16 km W of Alamos; 24 km W of Agiabampo; 32 km SW of Sonoyta (all AMNH). Sinaloa: Elota (AMNH). Nuevo Leon: Linares (AMNH). Tamaulipas: Mante (AMNH). San Luis Potosi: 1.6 km SW of Tamazunchale; Valles (both AMNH); 3 km S of Valles (MCZ). Nayarit: Acaponeta; Rio Las Canyas, 5 km NW Acaponeta; Jalisco; Mecatan; Rosamorada; 42 km S of Rosamor- ada; Tepic (all AMNH). Jalisco: near Anonas (AMNH). Hidalgo: Chapulhuachn (AMNH); Tzindejeh (CAS). Veracruz: Catameco (Playa Azul); Fortin; Veracruz (all AMNH). Michoacan: Jiquilpan; 4.8 km W of Quiroga (both AMNH). Guerrero: 1 1 km S of Chil-



================================================================================

Figures 1-5, Larinia direcia (Hentz). 1-4, female. I, epigynum, ventral. 2, epigy- num, posterior. 3, dorsal. 4, abdomen, ventral. 5, male, left palpus, mesal. Figures 6-9, Larink lampa n. sp., female. 6, epigynum, ventral. 7, epigynum, posterior. 8, dorsal. 9, abdomen, ventral. Figures 10- 15, Larinia fucuman n. sp. 10- 13, female. 10, epigynum, ventral 1 1, epigynurn, posterior. 12, dorsal. 13, abdomen, ventral. 14, 15, male. 14, palpus, mesai, 15, palpus, ventral.
Scale lines, 1.0 mm, genitalia 0. I mm. Abbreviations. A, terminal apophysis; C, conductor; E, embolus; M, median apophysis; T, tegulum.



================================================================================

252 Psyche [VOI. 97
pancingo (AMNH). Oaxaca: Oaxaca (AMNH). Chiapas: 105 km SE of Palenque (MCZ). Quintana Roo: Chetumal (MCZ). HON- DURAS Morazan: 27 km S of Tegucigalpa (MCZ). Atldntida: Lancetilla (MCZ). NICARAGUA Corn Island (MCZ); Masawas, Huaspuc River (AMNH); Islas de Solentiname (MAES); Leon (M AES). COSTA RICA Cartago: Turrialba (CAS, MCZ). Here- dia: Puerto Vieja (MCZ). Puntarenas: Monteverde Community (MCZ). PANAMA Bocas del Toro: (MIUP). Canal Zone: Fort Kobbe (AMNH). Chiriqui: Fortuna (MIUP). Panama: Soberania Natl. Park (MIUP). BAHAMAS common (AMNH). CUBA Real Cambina: Aguada, Las Villas (AMNH). Santa Clara: Topes de Col- lantes (AMNH). Soledad: Cienfuegos (MCZ). JAMAICA Grove Place; Long Mt.; Lucea (all AMNH). COLOMBIA Antioquia: La Estrella; Medellin (both MCZ). Cesar: Curumani (AMNH). Mag- dalena: Santa Marts (SMF), Meta: Carimagua (MCZ). Valle: Rio Tulua; Cali (both MCZ). SURINAM Matapica Beach (AMNH). VENEZUELA Zulia: Rosario (AMNH). ECUADOR Los Ri'os: Zapotal (CAS). PERU Madre de Dios: Alto Madre de Dios (MHNSM). BRAZIL Rio Branco: Roraima (MZSP). Amapa: San- tana (MEG). Amazonas: Ilha da Marchantario-Rio SolimGes (MCN). SZo Paulo: Guarujh (AMNH). PARAGUAY Alto Parana: Taquarazapa ( AMNH).
Larinia lampa new species