Diversidade de artrópodes nos abrigos foliares produzidos por Gonioterma sp. (Lepidoptera) em ramos de Roupala montana Aubl. (Proteaceae) no Cerrado do Brasil central

Gonioterma sp. é uma espécie monófaga de lepidóptero que constrói seus abrigos foliares (AFs) em Roupala montana, uma espécie arbórea comum na vegetação dos cerrados do Brasil central. No Parque Estadual de Caldas Novas (PESCAN), estado de Goiás, foi realizado um estudo que procurou investigar: 1) preferência do lepidóptero em construir seu AF de acordo com o tamanho da planta; 2) a composição, riqueza e as guildas dos artrópodes associados aos AFs; e, 3) possível associação entre o tamanho das plantas com a abundância dos artrópodes presentes nesses AFs. Os principais resultados revelaram que as mariposas fêmeas de Gonioterma sp. podem depositar seus ovos em R. montana com diferentes classes de tamanhos, porém com preferência por plantas de maior porte. Os AFs apresentaram uma elevada riqueza e abundância de artrópodes de diferentes guildas, onde os mais representativos foram os predadores. Foi verificado associação positiva entre o tamanho de R. montana e a ocorrência de determinadas guildas de artrópodes. O papel exercido por Gonioterma sp. como engenheiro de ecossistema ficou destacado pela importância dos abrigos foliares produzidos por este animal como refúgios para a fauna de artrópodes locais

Ecology and evolution of social behavior in Neotropical pseudoscorpions: the example of Paratemnoides nidificator (Atemnidae).

In the last two decades new species were discovered living in complex social organizations, besides the hymenopterans and isopterans. Examples as the naked mole-rat, coral-reef shrimps, aphids, thrips and beetles were added to the lists of eusocial species. Intermediate degrees of sociality were also described in arachnids as the spiders, amblypygids and uropygids, harvestmen, scorpions, and now pseudoscorpions. Although there is great resistance in the use of social behavior classifications, which usually privilege the eusocial species, we cannot deny that a lot of species took convergent pathways. Independently of the degree of social complexity, each species has unique perspectives in understanding the evolution of cooperative behaviors, especially the intermediate species. In this manuscript I present the natural history of a small social-permanent arachnid, capable to live in large colonies maintained by collective work and complex cooperative behaviors. Although it is not a new species for the science, we know very little about it social behavior. Now, we know that only two among the more than three thousand known pseudoscorpion species live in complex obligate societies. In this study we will present the natural history of Paratemnoides nidificator (Balzan, 1888) (Atemnidae) and their differences in relation to solitary pseudoscorpions; cooperative forage and dispersion and it implications for the maintenance of sociality in this group. We will discuss the existence and the evolution of division of labor; and also the existence that a second way of social life based on parasitism; and finally, a revision about the social behavior in the Pseudoscorpiones order and an evaluation of the main factors in the selection this way of life. These small and discreet animals can tell us a surprising history and help us to better understand the evolution of the social behavior in arthropods.Doutor em Ecologia e Conservação de Recursos NaturaisNas últimas duas décadas foram descobertas novas espécies vivendo em complexas organizações sociais, além dos himenópteros e isópteros. Exemplos como ratos-toupeira, camarões, afídeos, pulgões e besouros foram adicionados às listas de espécies eussociais. Estágios intermediários de socialidade também foram descritos em novos aracnídeos como as aranhas, amblipígeos e uropigídeos, opiliões, escorpiões, e agora pseudoescorpiões. Embora ainda existam grandes ressalvas no uso das classificações do comportamento social, que geralmente privilegiam as espécies eussociais, não se pode negar que muitas espécies tomaram caminhos convergentes. Independentemente do grau de complexidade social, cada espécie tem um poder único de abrir novas perspectivas na compreensão da evolução dos comportamentos cooperativos, em especial as espécies intermediárias. Neste manuscrito apresento a história de um pequeno aracnídeo social obrigatório, capaz de constituir grandes colônias mantidas pelo trabalho coletivo e complexos comportamentos cooperativos. Embora não seja novo para a ciência, ainda conhecemos muito pouco sobre seu comportamento social. Atualmente, sabemos que apenas duas dentre as mais de três mil espécies conhecidas de pseudoescorpiões vivem em complexas sociedades permanentes. Neste volume apresentaremos a história natural de Paratemnoides nidificator (Balzan, 1888) (Atemnidae) e suas diferenças em relação aos pseudoescorpiões solitários, comportamento cooperativo de forrageio e de dispersão e sua implicação para a manutenção da socialidade neste grupo; discutiremos a existência e a evolução da divisão de trabalho; também a existência que um segundo modo de vida social baseada em parasitismo em outra família; por fim, uma revisão sobre o comportamento social na ordem pseudoescorpiões e uma avaliação dos principais fatores na seleção deste modo de vida. Estes pequenos e discretos animais podem nos contar uma surpreendente história, e nos ajudar a compreender melhor a evolução do comportamento social em artrópodes

Natural history and behavior of the social pseudoscorpion Paratemnoides nidificator (Balzan, 1888) (Arachnida): parental care, cooperation and division of labor

The social behavior is rare among the arachnids. The most complex forms are found in the cooperative spiders, characterized by generation overlap, communal nests, food sharing and cooperative parental care. The two unique social pseudoscorpion species are found in South America. These species present a very similar social structure observed in the cooperative spiders, denominated "non-territorial permanent sociality". The present study investigated the natural history and the social behavior of Paratemnoides nidificator, a very common social Atemnidae in the Brazilian Savannah (Cerrado), at field and laboratory, over four years. This pseudoescorpião constitutes large family colonies, characterized by cooperative parental care extended until that the nymphs reach the adult phase, which can culminate in matriphagy. P. nidificator is a sit-and-wait generalist predator, capable to attack and subdue cooperatively preys sixty times heavier than a single individual. The dispersion in this species occurs by two ways: phoresy and colony fission. The Paratemnoides colonies present etarian and sexual division of work, in which individuals of different sex or stage can specialize in specific tasks colony maintenance. This is the first wide study about the natural history and behavior of a social pseudoscorpion. All these information, together, allow a wide comparison of the pseudoscorpions and spiders social organization.Mestre em Ecologia e Conservação de Recursos NaturaisO comportamento social é raro entre os aracnídeos. As formas mais complexas são encontradas nas aranhas cooperativas, existindo sobreposição de gerações, ninhos comunais, partilha de alimento e cuidado parental cooperativo. As duas únicas espécies sociais de pseudoescorpiões são encontradas na América do Sul. Estas espécies apresentam a mesma estrutura social observada nas aranhas cooperativas, denominada socialidade permanente não-territorial . O presente estudo investigou a história natural e o comportamento social de Paratemnoides nidificator, um atemnídeo social muito comum no Cerrado brasileiro, em campo e laboratório ao longo de quatro anos. Este pseudoescorpião constitui grandes agradados familiares, caracterizado por cuidado parental cooperativo estendido até que os filhotes atinjam a fase adulta, que pode culminar em matrifagia. P. nidificator é um predador de espreita generalista, capaz de atacar e abater cooperativamente presas sessenta vezes mais pesadas que um único indivíduo. A dispersão nesta espécie ocorre de duas formas: por forésia e fissão da colônia. Os agregados são caracterizados por divisão etária e sexual de trabalho, na qual indivíduos de sexo ou estádio diferentes se especializam em tarefas específicas de manutenção da colônia. Este é o primeiro estudo tão abrangente sobre a história natural e comportamento de um pseudoescorpião social. Estas informações, em conjunto, permitem uma ampla comparação da organização social de pseudoescorpiões e aranhas cooperativas

MATRIPHAGY IN THE NEOTROPICAL PSEUDOSCORPION PARATEMNOIDES NIDIFICATOR (BALZAN 1888) (ATEMNIDAE)

Volume: 33Start Page: 873End Page: 87

Report of Sphenochernes camponoti (Beier, 1970) (Pseudoscorpiones, Chernetidae) in phoresy on Fanniidae (Diptera)

Phoresy is a common dispersal behavior among pseudoscorpions. Neotropical pseudoscorpions, mainly from the North and Northeast regions of Brazil, are known for their dispersal relationships with beetles and flies. Here, we report phoretic association among nymphs of Sphenochernes camponoti (Chernetidae) and Fannia flies (F. pusio, F. yenhedi, and F. canicularis) (Diptera, Fanniidae). Twelve flies, each carrying a young pseudoscorpion, were collected in Caatinga vegetation in Pernambuco State, Brazil. Sphenochernes camponoti is a myrmecophilous pseudoscorpion that lives in Camponotus and Acromyrmex colonies. Despite its association with ants, this pseudoscorpion uses other winged arthropods to disperse. This is the first report of phoresy by Sphenochernes camponoti.

New records of pseudoscorpions (Arachnida, Pseudoscorpiones) from the Caatinga biome, Brazil: a checklist and a map of species richness distribution

In the course of ongoing research on the pseudoscorpion fauna in the northeastern region of Brazil, we compiled nine pseudoscorpion species with three of them, Geogarypus amazonicus, Apolpium ecuadorense, and Pachyolpium furculiferum, recorded for the first time from the Caatinga biome. Ecological comments are included, and the presence of G. amazonicus is discussed. Additionally, an updated checklist of all pseudoscorpion species and a map of the distribution of species richness for all biomes of Brazil are presented

Manuscript data

Raw data from manuscrip

Ceriochernes foliaceosetosus Beier 1974

Ceriochernes foliaceosetosus Beier, 1974 Material examined. 1 female, Pernambuco, Brazil: Meu Rei cave, 08°29’14.1”S, 37° 16’48.8”W, 777 m a.s.l., 19- I-2016, Catimbau National Park, E. Barbier leg. (LECA; Ps-012). Diagnosis. According to the description made by Beier (1974), the specimens of Ceriochernes foliaceosetosus Beier, 1974 can be easily distinguished by the carapace reddish-brown, slightly longer than wide, without eyes and eyespots, strongly granulated, the granules of the metazone covered with an epicuticle, reticulate in the form of a rough honeycomb, both transverse furrow narrow and not very deep, but distinct, the anterior curved laterally, the posterior closer to the rear margin, with six setae on the anterior margin, and eight on the posterior margin. All tergites divided, setae clavate and broadly foliated, rounded at the tip, curved and almost sessile, partially covered by epicuticle, with midrib, pleural membrane very densely granulated. Chaetotaxy tergal: 6: 8: 8: 6: 6: 8: 8: 8: 8: 8: 8: 2. Genital region operculum with a set of 12 setae, and ten setae on posterior margin. Chelicerae with six to seven setae on hand, es very short, three blades on the rallum, Serrula with 17 blades, galea with 5 very short branches distally. Pedipalps strongly reticulated, trochanter 1.0×, femur 2.58×, patella 2.15×, fingers not as long as the hand without pedicel, each with about 35–40 marginal teeth, the accessory teeth small and inconspicuous, present on the medial side only on the distal part of the finger, trichobothrium ist midway between isb and it, est proximal of ist; trichobothrium st closest to t. Measurements (mm). Body length: 2.021. Carapace: 0.638 / 0.538. Pedipalps: trochanter: 0.344 / 0.244, femur: 0.522 / 0.204, patella: 0.569 / 0.204, chela with pedicel: 0.864 / 0.346, chela without pedicel: 0.833 / 0.346, movable finger: 0.383. Leg I: trochanter: 0.124 / 0.105, femur: 0.127 / 0.093, patella: 0.255 / 0.093, tibia: 0.275 / 0.072, tarsus: 0.283 / 0.053. Leg IV: trochanter: 0.183 / 0.140, femur+patella: 0.488 / 0.152, tibia: 0.377 / 0.109, tarsus: 0.385 / 0.088. Distribution. Brazil (Harvey 2013; World Pseudoscorpiones Catalog 2022).Published as part of Bedoya-Roqueme, Edwin, Tizo-Pedroso, Everton, Barbier, Eder & Lira, André Felipe De Araújo, 2023, Two new cave-dwelling pseudoscorpion species (Arachnida: Pseudoscorpiones) from Northeastern Brazil, pp. 317-332 in Zootaxa 5293 (2) on page 321, DOI: 10.11646/zootaxa.5293.2.6, http://zenodo.org/record/796023

Progarypus smaugi Bedoya-Roqueme & Tizo-Pedroso & Barbier & Lira 2023, sp. nov.

Progarypus smaugi sp. nov. Figs. 3 – 4, 15 – 24 urn:lsid:zoobank.org:act:003a5f97-862a-4dee-94d1-98722cbad9ff Material examined. Male holotype: Pernambuco, Brazil: Meu Rei cave, 08°29’14.1”S, 37°16’48.8”W, 777 m a.s.l., 19-I-2016, Catimbau National Park, E. Barbier leg. (LECA; Ps-016). Paratypes: Brazil: 3 males, Pernambuco, Meu Rei cave, 08°29’14.1”S, 37°16’48.8”W, 777 m a.s.l., 19-I-2016, Catimbau National Park, E. Barbier leg. (LECA; Ps-017); 4 females, Pernambuco, Meu Rei cave, 08°29’14.1”S, 37°16’48.8”W, 777 m a.s.l., 19-I-2016, Catimbau National Park, E. Barbier leg. (LECA; Ps-018). Diagnosis. Distribution of the trichobothria on the fixed finger chelal, being trichobothrium ist distal de est. Trichobothrium it proximal to est; ib near the level of isb. Nodus ramosus on movable finger chelal proximal to trichobothria t; the robust pedipalps, male femur 0.604 / 0.229 mm, 2.6× (female, 0.681 / 0.251 mm, 2.7×), patella 0.485 / 0.209 mm, 2.3× (female, 0.552 / 0.250 mm, 2.5×). Fixed finger with 39 pointed teeth, movable finger with 30 flattened teeth. Adults. Carapace (Figs. 3–4), and pedipalps reddish-brown, carapace in both male and female with pale posterior area, more evident in males than in females, all legs light yellow, all tergites divided (except XI), anterior tergites light-brown colors, chela pedipalpal darker reddish-brown (Figs. 3–4). Chelicera with five setae on hand, fixed finger with one apical tooth, and four subapical teeth, movable finger with tooth-like subapical lobe (Fig. 16), rallum with three blade, first and second long toothed (Fig. 17), gale long, with three apical branches (Fig. 18), serrula exterior with 18 blades (female, with 20 blades). Pedipalps robust (Figs. 19–20), in both male and female strongly granulated, trochanter 1.3× (female, 1.2×) as long as broad, femur 2.6× (female 2.7×) as long as broad, chela with pedicel 3.2× (female 3.4×), chela without pedicel 3.0× (female 3.2×), inner margin of hand in female rounded (Fig. 20). Movable finger chelal 0.518 mm (female 0.641 mm), fixed finger with 39 pointed teeth (female with 37 pointed teeth), movable finger with 30 flattened teeth (female with 32 flattened teeth) (Fig. 21). Trichobothrium ist distal de est; trichobothrium it proximal to est; ib near the level of isb; trichobothrium st proximal to the middle of the finger, sb closer to b than to st, nodus ramosus on fixed finger distal of est, and in the movable finger chelal proximal to trichobothrium t (Fig. 21), well-developed venom apparatus in both on the fixed finger and on the movable finger chelal (Fig. 21). Carapace triangular (Fig. 15), squamose sculpturing, posteriorly reticulate, cucullus shorts, with an indistinct longitudinal furrow, with medial furrow and indistinct posterior furrow (Fig. 15), four large eyes, all setae very short and acuminate, with approx. 38 setae (female approx. 36 setae), four setae on the anterior margin, one seta preocular on each side, with six to twelve setae on the posterior margin (Fig. 15). Coxal region: female and male with an apical lobe of coxa pedipalpal with two setae marginal, and three discal setae, approx. 26 setae (female approx. 28), coxa I ca. 22 setae (female ca. 26 setae), Coxa II ca. 24 setae (female ca. 28), Coxa III ca. 30 setae (female ca. 32 setae), Coxa IV ca. 34 setae (female ca. 36 setae). Opisthosoma: all tergites are divided (except XI); reticulate sculpture, all setae acuminate, pleural membrane strongly striated. Tergal chaetotaxy of holotype male, tergites I–XI: 6: 8: 8: 8: 6: 6: 4: 4: 6: 6: 4 [4ST]: 2, females paratypes, tergites I–XI: 6: 6–8: 6–8: 8: 6: 6: 6–4: 6–4: 6: 6: 4 [4ST]: 2, males paratypes, tergites I–XI: 6: 6: 8: 8: 6: 6–8: 6–8: 6–8: 6: 6–8: 4 [4ST]: 2, tergite XI with four setae tactile. Sternal chaetotaxy of the male holotype, sternites II–XII: 8(4): (1)6(1): (1)6(1): 8: 8: 8: 8: 8: 10: 10: 2, females paratypes, sternites II–XII: 6: (1)6(1): (1)6(1): 6–8: 6–8: 8: 8–10: 8–10: 10–12: 10: 2, males paratypes, sternites II–XII: 8(4): (1)6(1): (1)6(1): 6: 6–8: 8–10: 8–10: 8–10: 10–12: 10: 2, sternites XI with two setae tactile, pleural membrane III–X with one seta. The genital region of the male holotype with eighth setae, and four discal setae on the anterior operculum (Fig. 22), the posterior operculum with six marginal setae (Fig. 22), internal genital structures, indistinctly visible (Fig. 22). Genital region of the female paratype with six central setae on the anterior operculum, and six marginal setae on the posterior operculum. Legs: all legs light-yellow (Figs. 23–24), Leg I (Fig. 23): trochanter 1.5× (female 1.7×), femur 2.8× (female 2.7×), patella 2.4× (female 2.6×), tibia 2.4× (female 2.6×), metatarsus 3.2× (female 3.3×), tarsus 3.3× (female 4.8×), Leg IV (Fig. 24): trochanter 1.1× (female 1.2×), femur+patella 2.6× (female 2.7×), tibia 4.0× (female 4.1×), metatarsus 3.1× (female 3.2×), tarsus 3.7× (female 3.8×), subterminal setae smooth, arolium undivided and longer than claws (Fig. 24). Measurements (mm):Male holotype (Paratype in parentheses):body length 1.796(1.813); carapace 0.609 / 0.486 (0.612 / 0.489); Chelicerae; movable finger. pedipalps trochanter: 0.255 / 0.191 (0.257 / 0.193); femur: 0.604 / 0.229 (0.607 / 0.230); patella 0.485 / 0.209 (0.487 / 0.210); chela with pedicel 0.929 / 0.289 (0.931 / 0.290); chela without pedicel 0.873 / 0.289 (0.853 / 0.290), movable finger 0.578 (0.581). Leg I: trochanter 0.136 / 0.129 (0.137 / 0.131), femur 0.220 / 0.077 (0.225 / 0.079), patella 0.149 / 0.061 (0.151 / 0.063), tibia 0.159 / 0.065 (0.161 / 0.067); metatarsus 0.131 / 0.041 (0.133 / 0.043); tarsus 0.134 / 0.043 (0.136 / 0.045). Leg IV: trochanter 0.231 / 0.131 (0.235 / 0.133); femur+patella: 0.843 / 0.180 (0.845 / 0.183); tibia: 0.361 / 0.091 (0.365 / 0.093); metatarsus 0.219 / 0.071 (0.221 / 0.070); tarsus 0.189 / 0.051 (0.187 / 0.053). Female paratypes: Body length 2.241 –2.250. Carapace 0.691–0.693/0.552–0.553; Chelicerae; movable finger. Pedipalps:trochanter 0.226–0.229/0.210–0.213; femur 0.681–0.683/0.251–0.253; patella 0.550–0.553/0.220–0.223; chela with pedicel1.091–1.093/0.321–0.323; chela without pedicel 1.031–1.033/0.321–0.323; movable finger 0.641 – 0.643. Leg I: trochanter 0.050–0.060/0.136–0.137; femur 0.263–0.265/0.080–0.090; patella 0.167–0.169/0.102– 0.103; tibia 0.215–0.217/0.080–0.090; metatarsus 0.203–0.206/0.060–0.070; tarsus 0.195–0.197/0.041–0.043. Leg IV: trochanter 0.230–0.231/0.130–0.133; femur+patella 0.803–0.805/0.251–0.253; tibia 0.532–0.533/0.110–0.112; metatarsus 0.316–0.319/0.060–0.070; tarsus 0.249–0.251/0.040–0.050. Etymology. The specific epithet (smaugi) derives its name from the mythical (fictional) dragon, the last of the great dragons remaining in Middle Earth, in the fantasy literary book “The Hobbit” written by J.R.R. Tolkien. Distribution. Only known from the type locality (Brazil: Pernambuco, Meu Rei cave). Remarks. According to the descriptions by Beier (1931a, 1932b, 1959, 1964) and the taxonomic key presented by Mahnert (2001), Progarypus smaugi sp. nov. differs from other species of Progarypus Beier, 1931 as follows: it differs from the type species of the genus Progarypus ramicola (Balzan, 1887), from Paraguay, for the femur pedipalpal without pedicel, clearly separated, somewhat concave, femur pedipalps 0.720 / 0.160, patella 0.690 (0.540)/0.201, rallum of chelicera with three denticulate blades. It differs from Progarypus longipes Beier, 1964 by each tergite with six to eight very short marginal setae; pedipalps relatively very thinned, femur pedipalpal on the apical third with an extended tactile seta, scaly sculpture in the medial area and granular in the rest. Fixed finger chelal with 45 densely spaced teeth, finger movable chelal with 35 pointed teeth in apical third, but completely flat at the base, and Progarypus marginatus Beier, 1964 from Chile by the eb -esb -isb trichobothria at the base of the fingers, ist is closer to it than to ib; male femur pedipalpal 0.950 / 0.201, 4.7×, patella 0.801 / 0.240, 3.3×. Differ from Progarypus novus Beier, 1931 from Brazil by the carapace 1.5× as long as wide, gradually narrowing, with truncated anterior margin, and approximately net-like sculpturing. Pedipalps slender, femur 4.0×, as long as wide, sessile, patella 3.5×, chelal fingers slightly shorter than hand without pedicel 0.54 mm. It differs from Progarypus peruanus Beier, 1959 from Peru by the carapace as long as wide at the base, conically narrowed at anterior margin, densely granulated. Pedipalps almost as long as the body, femur without pedicel 4.3×, patella 3×, fingers clearly longer than hand chela with pedicel and as long as femur, tarsus IV shorter than tarsus I. It differs from Progarypus oxydactylus (Balzan, 1887) from Paraguay by the carapace clearly longer than wide, finely granulated, and reticulate at the base, pedipalps rather long and thin. Femur 4.1× times as long as wide, weakly pedicelled, finely granulated, and striated, patella 3.2× as long as wide, notably shorter than the femur. Similarly, it differs from the species Progarypus gracilis Mahnert, 2001, Progarypus liliae Mahnert, 2001, and Progarypus setifer Mahnert, 2001 from Brazil by the length of the femur pedipalps 0.7-1.0- 1.21 mm, by the distribution of the trichobothria est, ist, it, et, the length of the fingers of the pedipalpal hand. The new species resembles Progarypus nigrimanus Mahnert, 2001, from Brazil however, it differs by the femur pedipalpal length 0.7–1.0 mm; at least one of trichobothrium est, ist, it, et proximal to st, finger relatively shorter, at most 1.3×, pedipalps bicolored, chelal hand darker than femur and patella, femur pedipalpal 4.7–5.3× (length 0.86 mm), patella 3.8–3.9× (length 0.78–0.79mm) longer than broad, carapace anteriorly with tessellated sculpturing.Published as part of Bedoya-Roqueme, Edwin, Tizo-Pedroso, Everton, Barbier, Eder & Lira, André Felipe De Araújo, 2023, Two new cave-dwelling pseudoscorpion species (Arachnida: Pseudoscorpiones) from Northeastern Brazil, pp. 317-332 in Zootaxa 5293 (2) on pages 325-327, DOI: 10.11646/zootaxa.5293.2.6, http://zenodo.org/record/796023