Bioacoustics of Ephebopus cyanognathus

Characters : In characters marked with an asterisk (*; viz., 6, 10, 12, 13, 18, 20, 31, 39, 42), fit and steps are not given as the character is not significant to the analysis but was kept here to indicate an autapomorphy for a terminal taxon. (0) Eye tubercle in males and females: (0) low ( Fig. 23 ), (1) high ( Fig. 24 ); fit = 66.6. Steps/ Extra steps = 4/3. The eye tubercle can be absent in some mygalomorphs (e.g., Atypidae, Rastelloidina, Raven 1985 ; Goloboff 1993b , 1995 ) or well-defined in most mygalomorph taxa. Intermediate states have been proposed for some cyrtaucheniids, hexathelids, ctenizids ( Raven 1985 :125, 70, 141; Bond & Opell 2002 ), and diplurids ( Ischnothele Ausserer ) ( Bond & Opell 2002 ). In theraphosids, two states have been proposed: absent, for some Hemirrhagus Simon 1903 , ( Spelopelma Gertsch ; Raven 1985 ) and distinct, for remaining species. An intermediate state was herein identified: Heteroscodra ( Fig. 23 ) shows a somewhat flattened eye tubercle, with anterior lateral eyes being in almost same plane as anterior median eyes (state 0). Spiders with an arched eye tubercle ( Fig. 24 ), with the anterior median eye positioned clearly in superior plane in relation to the anterior lateral eye were considered as having state 1. (1) Anterior row of eyes in males and females, measured from anterior margins: (0) procurved, (1) straight; fit = 85.7. Steps/ Extra steps = 2/1. (2) Clypeus in males and females: (0) absent, (1) narrow ( Fig. 23 ) (2) wide ( Fig. 24 ); fit = 60.0. Steps/ Extra steps = 6/4. The clypeus was considered absent when there is no space between the eye tubercle and the anterior carapace edge which can slightly project forward beyond the anterior carapace edge. A wide clypeus is found in some theraphosids, e.g. Harpactirinae , Ornithoctoninae and some theraphosines. In this case, the distance between the anterior edge of the eye tubercle and the anterior edge of carapace can be almost the same as the length of the eye tubercle ( Fig. 24 ). The narrow state of the clypeus occurs in intermediate cases ( Fig. 23 ). (3) Fovea in males and females, curvature: (0) straight, (1) recurved, (2) procurved; fit = 85.7. Steps/ Extra steps = 3/1. (4) Fovea in males and females, closure: (0) slit-like (closed), (1) pit-like (open); fit = 100. Steps/ Extra steps = 1/0. In Mygalomorphae, a pit-like (open) fovea seems to be plesiomorphic, since Mesothelae shows this state as well as representatives of Atypidae , Antrodiaetidae and Dipluridae ( Raven 1985 ) . For theraphosids, Gallon (2003) suggested the presence of pit-like fovea to be a synapomorphy of the clade Xenodendrophila Gallon 2003 (now Encyocratella Strand ) ( Stromatopelma + Heteroscodra ). This proposal is followed here. (5) Fovea in males and females, depth: (0) shallow; (1) deep; fit = 66.6. Steps/ Extra steps = 4/3. This character is introduced here and refers to the depth of the fovea. Plaster was applied to the specimen's fovea and the resulting mould was analyzed. Those with deep fovea (state 1) presented a mould with trapezoid shape. Specimens with shallow fovea (state 0) showed a mould with a short line. (6) Labial cuspules in males and females, number*: (0) 30–300, (1) 0–20, (2) 350–450 Within the Theraphosidae examined here, only Euathlus has a low number of labial cuspules. The most common state is between 30-300 labial cuspules and the only taxon with more than this number is Phlogiellus . The state 30– 300 was coded as 0 because it is present in the outgroups and thus should be the plesiomorphic state. (7) Sigilla, posterior pair in males and females, position: (0) marginal, less than 1.5 times its own diameter from margin, (1) located closer to center of sternum separated from margin by twice its own diameter; fit = 75. Steps/ Extra steps = 3/2. In Eumenophorinae, the posterior sternal sigilla are medially placed ( Pocock 1897 ; Gallon 2003 ). We consider Phlogiellus and Haplopelma also have this state. (8) Tarsus IV in males, cracked: (0) cracked, (1) integral; fit = 85.7. Steps/ Extra steps = 2/1. A cracked tarsus is found in some mygalomorph taxa, e.g., some diplurids, nemesiids, barychelids and theraphosids ( Raven 1985 ). In specimens examined, this condition was found in males and females of Melloina and males of Holothele and Phlogiellus . (9) Tarsal scopulae in males and females: (0) no true scopula; (1) scopula of sparse hairs; (2) dense scopula that does not extend much laterally; (3) scopula very extensive laterally, giving the tarsi and metatarsi I and II a spatulate appearance; fit = 85.7. Steps/ Extra steps = 4/1. Melloina has no true scopula (state 0) ( Raven 1999 ). In Holothele and other Ischnocolinae, the scopula is very sparse (state 1), and frequently divided by rows of setae. Most theraphosids have well-developed scopulae (state 2), that, however, is not so developed as those having state 3, where their lateral development give the tarsi and the metatarsi I and II a spatulate appearance. This last state was used by Simon (1892) to characterize the Avicularieae and that was followed by most authors thereafter. (10) Scopulae on metatarsus IV in males and females*: (0) divided by hairs or spines; (1) not divided. Gallon (2003) considered the undivided scopula on metatarsus IV to be apomorphic for Eumenophorinae. The only taxon with the derived state in the analysis is Citharischius Pocock 1901 . (11) Leg spines in males and females: (0) present on apex and other faces of tibiae and metatarsi, (1) present only on ventral apices of tibiae and metatarsi, (2) absent; fit = 75. Steps/ Extra steps = 4/2. The theraphosid sister-groups, e. g., Melloina and barychelids, have spines distributed largely on ventral, prolateral, retrolateral and sometimes dorsal faces of the legs. This suggests that the condition is plesiomorphic for Theraphosidae . Among the studied taxa, a lower number of leg spines is seen in Tapinauchenius , Psalmopoeus , Ephebopus , Citharischius and Phlogiellus where there are only two small spines on the ventral apical edge of the tibiae and metatarsi. In Avicularia , Iridopelma , Pachistopelma , Stromatopelma , Heteroscodra and Poecilotheria , spines are totally absent. (12) Palpal femora in males and females, scopula on retrolateral face*: (0) absent, (1) present. Raven (1985) included Stromatopelma in the Eumenophorinae because they share the presence of a brush of hair on the retrolateral face of the palpal femora. Gallon (2003) stated that only Stromatopelma females present this scopulae and proposed its presence to be homoplastic with respect to Eumenophorinae. We failed to find this structure in males and females of Stromatopelma examined, and so it was here coded as absent. The only genus in the analysis clearly having the scopulae was Citharischius . (13) Femora IV in males and females, scopulae on retrolateral face*: (0) absent, (1) present.The scopulae on the retrolateral face of femora IV is an apomorphy of Lasiodora C. L. Koch 1850 and other theraphosine genera not included in the analysis ( Pérez-Miles et al. 1996 ). (14) Chelicerae in males and females, scopulae on retrolateral face: (0) absent, (1) present; fit = 85.7. Steps/ Extra steps = 2/1. (15) Maxillae, spiniform setae on lower prolateral face: (0) absent, (1) present; fit = 100. Steps/ Extra steps = 1/0. Spiniform setae on the lower prolateral face of maxillae are found in Ornithoctoninae and Thrigmopoeinae ( Raven 1985 ). We here consider that spines in the same position in Poecilotheria are putative primary homologs. (16) Stridulatory bristles in males and females form maxillae lyra: (0) absent, (1) present; fit = 75. Steps/ Extra steps = 3/2. The derived condition (1) of this character was used to characterize the Selenocosmiinae (e.g., Pocock 1895 ). It is also found in other mygalomorph taxa apart from Theraphosidae , e.g., the diplurids Diplura C. L. Koch 1850 and Trechona C. L. Koch 1850 ( Raven 1985 ) . (17) Stridulatory bristles on coxae I of males and females: (0) absent, (1) present; fit = 85.7. Steps/ Extra steps = 2/1. Stridulatory bristles on coxae were used to characterize the Eumenophorinae (e.g., Raven 1985 ) and some Theraphosinae genera such as Lasiodora , Grammostola Simon 1892 and Theraphosa Thorell 1870 ( Pérez-Miles et al. 1996 ) . (18) Longitudinal white stripes on patellae and tibiae of males and females*: (0) absent, (1) present ( Fig. 11 ). The conspicuous white stripes in Ephebopus murinus were considered an autapomophy. (19) Leg rings on distal femora, tibiae and metatarsi of males and females: (0) absent, (1) white, (2), yellow ( Figs 13–14, 16 ); fit = 85.7. Steps/ Extra steps = 3/1. Most theraphosid species have a band of short setae around the distal femora, tibiae and metatarsi of all legs, more visible on legs I and II. Some taxa, e. g. Holothele , have no detectable rings, whereas in most theraphosids, they are white, if present. Some species of Ephebopus and Avicularia have these rings but are yellow in color. (20) Chelicerae of females, hair: (0) not iridescent, (1) iridescent ( Fig. 16 ). State 1 is an autapomophy of E . cyanognathus . (21) Black marking dorsally on tibiae, metatarsi and tarsi of males and females: (0) absent, (1) present; fit = 100. Steps/ Extra steps = 1/0. This character was introduced by Gallon (2003) and his proposal is followed here. (22) Female abdominal pattern: (0) all one color or irregularly mottled, (1) with pattern; fit = 60. Steps/ Extra steps = 5/4. Gallon (2003) considered that ornithoctonines, harpactirines and "stromatopelmines" ( Heteroscodra , Stromatopelma and Xenodendrophila , now Encyocratella ) to have abdomen dorsums marked with a herring-bone pattern. We follow that here and include also Poecilotheria and E . foliatus as well as Avicularia , Pachistopelma , Iridopelma , Tapinauchenius and Psalmopoeus Pocock 1895 in which genera, immatures have a dorsal abdominal pattern. (23) Spermathecae, number: (0) two, completely separated, (1), two, fused at base, (2) one, totally fused; fit = 100. Steps/ Extra steps = 2/0. (24) Spermathecal lobes: (0) absent, (1) present; fit = 66.6. Steps/ Extra steps = 4/3. (25) Cymbium: (0) without spiniform process between lobes, (1) with spiniform process between lobes; fit = 100. Steps/ Extra steps = 1/0. This character was proposed by Raven 1985 as a putative synapomorphy of Aviculariinae . (26) Prolateral cymbium lobe shape: (0) rounded ( Fig. 26 ), (1) subtriangular ( Fig. 25 ); fit = 85.7. Steps/ Extra steps = 2/1. (27) Subtegulum: (0) small, (1) large, extending down the bulb for half of the tegulum ( Raven 1985 ); fit = 85.7. Steps/ Extra steps = 2/1. A large subtegulum is considered a Theraphosinae synapomorphy ( Raven 1985 ; Pérez-Miles et al. 1996 ). This character state 1 is also present in Haplopelma Simon 1892 . (28) Embolus curvature: (0) straight ( Figs 1, 3, 5, 7, 9 ), (1) curved to the retrolateral side ( Fig. 25 ); fit = 66.6. Steps/ Extra steps = 4/3. (29) Embolus length: (0) 1.5-2.5 times longer than tegulum, (1) shorter than tegulum, (2) more than 3 times longer than tegulum; fit = 60. Steps/ Extra steps = 6/4. (30) Embolus distal width: (0) thin, less than 1/5 of tegulum height ( Fig. 25 ), (1) thick, more than 1/3 of tegulum height ( Fig. 26 ); fit = 85.7. Steps/ Extra steps =2/1 (31) Embolus shape*: (0) not flattened, (1) slightly flattened, (2) very flattened. The states follow Bertani (2000 ; 2001 ). Haplopelma was found to have a very flattened embolus, similar to some theraphosine species, such as Theraphosa , Megaphobema Pocock 1901 and Sericopelma Ausserer 1875 . (32) Prolateral inferior keel of bulb: (0) absent, (1) present; fit = 100. Steps/ Extra steps =1/0. Several mygalomorph taxa have strongly developed keels on the male palpal bulb, e.g., species of Pycnothelinae ( Nemesiidae ), Barychelinae ( Barychelidae ) and Theraphosinae ( Raven 1985 ) . In Theraphosidae , the presence of keels was considered a synapomorphy of Theraphosinae ( Raven 1985 ; Pérez-Miles et al. 1996 ). Bertani (2000) created a terminology for theraphosine male palpal bulb keels, which is followed here. However, other theraphosid taxa have strongly developed keels, such as species of Haplopelma and Poecilotheria . These keels are similar in position to some theraphosine species and thus they are being putatively considered primary homologs in the cladistic analysis. (33) Prolateral superior keel on embolus ( Bertani 2000 ): (0) absent, (1) present; fit = 100. Steps/ Extra steps =1/0. (See character 32). (34) Apical keel on embolus ( Bertani 2000 ): (0) absent, (1) present; fit = 85.7. Steps/ Extra steps = 2/1. (See character 32). (35) Retrolateral keel on embolus ( Bertani 2000 ): (0) absent, (1) present; fit = 85.7. Steps/ Extra steps = 2/ 1. (See character 32). (36) Male tibial spurs: (0) present, two-branched, (1) present, with apical megaspine, (2) present, with spiniform setae, (3) absent; fit = 66.6. Steps/ Extra steps = 6/3. Male tibial spurs are a very homoplasious character in Theraphosidae ( Bertani 2001 ) and most other mygalomorph families ( Raven 1985 ). However, the presence of two-branched spurs seems to be plesiomorphic in Theraphosidae since it is present in all barychelid subfamilies and in Melloina (Paratropididae) , the proposed sister-groups of Theraphosidae ( Raven 1985 ) . (37) Hairs on metatarsi and tibiae I–IV of males: (0) normal, (1) long hairs laterally projected, forming a brush; fit = 85.7. Steps/ Extra steps = 2/1. Males of Tapinauchenius , Psalmopoeus , Stromatopelma and Heteroscodra all share the presence of hairs projecting laterally. (38) Urticating hairs of males and females: (0) absent, (1) on abdomen ( Cooke et al. 1972 ), (2) on pedipalps ( Marshall & Uetz 1990 ) ( Fig. 19 ). Fit = 85.7. Steps/ Extra steps = 3/1. (39) Type I urticating hairs in males and females*: (0) absent, (1) present. (40) Type II urticating hairs in males and females: (0) absent, (1) present; fit = 100. Steps/ Extra steps = 1/ 0. (41) Type III urticating hairs in males and females: (0) absent, (1) present; fit = 100. Steps/ Extra steps = 1/0. Considered a synapomorphy of Theraphosinae ( Pérez-Miles et al. 1996 ). (42) Type IV urticating hairs in males and females*: (0) absent, (1) present. (43) Type V urticating hairs in males and females: (0) absent, (1) present ( Figs 19–20 ); fit = 100. Steps/ Extra steps = 1/0. Type V hair ( Fig. 20 ) is present in Ephebopus murinus ( Marshall 1990 ) and all known species of this genus. (44) Burrow entrance (fossorial) of female: (0) simple with little or no silk, (1) collar of silk bound with surrounding debris ( Fig. 15 ), (2) trumpet-shaped ( Fig. 11 ); fit = 85.7. Steps / Extra steps = 3/1. (45) Eggsac type : (0) mobile, (1) fixed hammock, (2) fixed flat; fit = 100. Steps / Extra steps = 2/0. Here we follow the classification of Gallon & Gabriel (2006) (46) Habitat: (0) evergreen forest, (1) deciduous forest, (2) desert-scrub; fit = 85.7. Steps/ Extra steps = 3/ 1. (47) Habits of female: (0) hide in surface layers of soil, (1) arboreal, (2) opportunistic ( Figs 17–18 ), (3) fossorial ( Fig. 11 ). Fit = 75. Steps / Extra steps = 5/2. State 0 was reported in paratropidids ( Raven 1999 ). Most theraphosids make a burrow (state 3) or hide in retreats built in trees using silk (most aviculariines and Poecilotheria ) (state 1). An opportunistic burrowing behavior (state 2) is found in Tapinauchenius , Psalmopoeus and Ephebopus rufescens (RCW and SDM pers. obs.). They build a silken retreat which is not always made high in trees or other elevated structures. Furthermore, they commonly use soil and/or rocky substrata.