Bioacoustics of Stenopelmatus

( Fig. 3C ) Commonly known as "boyish" or Jerusalen Crickets ( Weissman, 2005 ), are peculiar Orthoptera with fossorial habits, eyes narrowed, abdomen inflated and separated from the thorax by a small pedicel ( Cadena-Castañeda & Torres, 2013 ) are the closest group to rolling leaf crickets or gryllacridines ( Song et al ., 2015 ; Vandergast et al ., 2018). To date there are 20 valid pecies of Stenopelmatus Burmeister ; discontinuously distributed from Canada to Costa Rica ( Burmeister, 1838 ; Gorochov & Cadena-Castañeda, 2016 ; Cigliano et al. 2018). For Mexico there are registered S. lycosoides Walker , S. calcaratus Griffini and S. lessonae Griffini (no precise location); S. guatemalae Brunner von Wattenwyl ( Chiapas , but also registered to Guatemala and Nicaragua ). S. histrio Saussure and S. minor Saussure , ( Veracruz ); S. nietei Saussure , ( Veracruz , Hidalgo , Tamaulipas and Coahuila ); S. longispinus Brunner von Wattenwyl ( Sinaloa ) ; S. talpa Burmeister , (initially without precise locality but recently recorded by Gorochov & Cadena-Castañeda, 2016 , for the states of Oaxaca , State of Mexico and Veracruz ) and S. fuscus Haldeman , which is registered for the northeast Mexican territory. However, the photographs of S. fuscus found in reports in which the species is reported, such as those of Berumen et al . (2006) and Barrientos-Lozano et al . (2013) do not confirm? to the original description and most likely were erroneously identified, like the previous two publications, the record for this species in Mexico is doubtful ( Weissman, 2001 ; Gorochov & Cadena-Castañeda, 2016 ; Cigliano et al ., 2018). These individuals were collected from May to December. Females were trapped the most in carrion traps. The morphospecies studied here is very similar to S. minor Saussure by its small size and black color, according to Weissman (2001) . Numerous species await description by Dr. David Weissman, commenting only that only on the east coast of the United States there may be about 50 undescribed species morphologically similar. Species of Stenopelmatus live under the sand or soil (lowland species and semi-desert areas) but almost nothing is known about the species that inhabit the mountains, some individuals of this morphospecies were seen in galleries or decaying logs, are omnivorous habits and subjected to stress can become cannibalistic. At the request of Dr. David Weissman this species is not described.

In the large number of specimens that we examined from both ANSP and UMMZ, there are series of small, black individuals from numerous localities in Mexico , some undoubtedly representing undescribed taxa. But without DNA or drumming data, we decline to discuss any of them further. On the other hand, there are several series of small to large sized adult specimens, with good locality data, that appear morphologically distinct and we now list those. But in the absence of DNA, drumming information, or a diagnostic morphological character, we do not formally describe any of them. In any case, we implore our colleagues: if you want to describe a new species of JC, you should have at least three of these four basic items: (1) a reasonable series including adult males, (2) calling drum, because without this, accurately determining the distribution range of a taxon is almost impossible, (3) documentation of the presence or absence of jumping ability since such a trait is restricted to Stenopelmatus and there are certain subclades within Stenopelmatus where no taxa jump, and (4) DNA data. A bonus would be karyotype information, although considering only our preliminary data discussed herein, such doesn’t help separate Stenopelmatus from Ammopelmatus .

4. Mexico . Puebla . Puebla . 1922. KB Kyser. ANSP. One adult female ( Fig. 195A ). This large, stocky specimen is probably in Stenopelmatus based on rear tibia spines and protruding ovipositor. While resembling A. comanchus , from state of Durango , Villa Lerdo, in size, her outer, most distal rear leg tibial spines are clearly different (Weissman et al. in prep). How can a JC that is ~ 40 mm in length be known from only one specimen? 5. Mexico . Veracruz , Mt. Orizaba , 10-12000’, 24-vii-1938 , EH Taylor. Light colored legs ( Fig. 196A ), head, and pronotum, small-medium sized, 4 adult females, 1 late instar female. ANSP . 6. Mexico . Puebla . Mt. Citlaltepetl (aka Mt. Orizaba ) 17-vi to 14-vii-1949 , 13000-14000’, LW Swan , CAS , 1 adult male ( Fig. 197 ), 2 adult females, 2 small nymphs. Swan used topo maps (pers. comm. to DBW, July, 1994), while he walked around the entire mountain, so elevations should be accurate. Timberline ~13,500’, dominant plant was grass Calamagrostis eriantha . Specimens collected in sandy areas with volcanic soil. Highest elevation Stenopelmatus known. We have another very high elevation collection record: Mexico , Mexico , Tlamacas , ~12000’, on Mt. Popocatepetl , 1-viii-1969 . S. & J. Peck. 1 light colored mid instar, UMMZ . 7. Mexico . Veracruz . Metlac Canyon, S06-39. We found, microsympatric with S. sartorianus , a second species of jumping Stenopelmatus , this one an early instar, 12 mm in length, also inside a log, that was hyperactive, all black whose antennae with a white annulus, and with no wing pads, that we were unable to raise to adult. 8. El Salvador : Cerro Verde , 5-vi-1976 , C. & M. Goodnight , UMMZ . Metapán, Hacienda Monte Cristo 7000’ cloud forest, 6-vii-1957 , A. & P. Starrett & TM Uzzell, Jr. 1 mid-late male similar to S. talpa , UMMZ. These are the only 2 records we have seen from this country . What percentage of Stenopelmatus species have now been described? Given the restricted distributions of many taxa in Mexico and Central America, the lack of widespread collecting in Honduras , Guatemala , and Nicaragua coupled with the numerous collections of small, black individuals in the collections from ANSP and UMMZ, and the scarcity of good series of adults with calling drums in any collection, we guess that as little as 30% of the species diversity of Stenopelmatus could be known.

A. For the 16 named Stenopelmatus taxa for which we have first-hand experience ( Table 2 ), 10 species jump or hop as juveniles and adults. The 6 taxa that do not jump are all small (< 25 mm ). Of the 4 non-jumping taxa for which we have DNA, 3 of them ( S. perote , S. faulkneri , and S. durango ) are recovered together ( Fig. 10 ). The 4 th nonhopper, S. hondurasito , is recovered closest to S. ecuadorensis in the ASTRAL and mtDNA analysis, but we don’t know if the latter hops or not. In the mtDNA tree ( Fig 9 ), the clade with S. hondurasito is recovered closer to the other non-hopping taxa. Such hops/jumps can be from 6 to 15 cm in distance, and originate when both hind femurs are cocked and rapidly extended similar to the action seen when grasshoppers jump. In raising many thousands of Ammopelmatus over the years, we have never seen a single one jump. This trait is apparently the only character that is 100% diagnostic for Stenopelmatus , even if not all species do it. There is no relationship ( Fig. 12 ) between hind femur length and hind femur width, perhaps analogous to the results of Lawler et al. (2019) who found no relationship between femur length: mass ratio in a grasshopper. On the other hand, the best jumper was S. sartorianus , a species with long, narrow femurs and tibias. These physical modifications appear to be similar in a jumping South Africa cockroach ( Picker et al. 2012 ). B. Tree climbing was postulated by Weissman & Lightfoot (2007) for Stenopelmatus and subsequently confirmed by Gutiérrez-Rodríguez & Riverón (2018) . We have never seen arboreal behavior in Ammopelmatus . C. Antennae are constantly in motion in S. sartorianus , resembling that seen in an active pompilid wasp. D. Virgin adults of both sexes of all species of Ammopelmatus generally drum well in captivity. Even in reluctant drummers, documentation was eventually successful. In many Ammopelmatus taxa, the drums are easy to hear at distances of 10m . In contrast, Stenopelmatus virgin adults are generally reluctant, much softer drummers. In fact, we were unable to document drumming in 3 species ( S. mineraldelmonte , S. piceiventris , S. hondurasito ) of Stenopelmatus despite repeated efforts with several virgin individuals of each species, and we speculate that not all Stenopelmatus taxa may drum. We wonder if this behavioral difference could be analogous to the decreased calling song stridulation seen in many, similarly nocturnal, New World tropical katydids, possibly as a result of high bat predation ( Belwood & Morris 1987 )? E. While a number of Ammopelmatus taxa possess a sex clarification drum ( Weissman 2001b ), such is only known in one species of Stenopelmatus - S. perote (see p. 64).