Selizitapia gen. nov. (Hemiptera: Fulgoromorpha: Flatidae) from tapia woodlands of Madagascar

A new monotypic genus of flatid planthoppers (Hemiptera: Fulgoromorpha: Flatidae), Selizitapia gen. nov., is described for Selizitapia pennyi gen. et sp. nov. (type species) from the island of Madagascar. Habitus, male and female external and internal genital structures of the new species are illustrated and compared with similar taxa. Selizitapia pennyi gen. et sp. nov. is endemic to Madagascar where it is known to date only from one locality in the Central Plateau and is associated with tapia woodland formation.


Introduction
On a global scale, Madagascar represents one of the world's top twelve 'mega diversity' hotspots with high levels of taxonomic endemism and species-rich tropical environments, being recognized as a naturalist's paradise (Myers et al. 2000;Goodman & Benstead 2005).
Especially the Malagasy entomofauna is extremely rich and many new species are added to the island check-list year by year (e.g., Chłond et al. 2018;Gnezdilov 2019;Kim et al. 2020).
However, despite major conservation efforts, many unique ecosystems are now in a very large extent fragmented and degraded and examples here can be the eastern littoral forest, the western dry deciduous forest and the central highland tapia woody formation, to mention only these three (Ganzhorn et al. 2001).
The Flatidae Spinola, 1839 are the fourth largest family of the infraorder Fulgoromorpha Evans, 1946, with 296 genera and 1443 species worldwide, respectively representing 12.0% and 10.5% of all planthoppers described (Bourgoin 2020). They are the most diverse in all regions of the tropics, far less diverse in the Holarctic (Bartlett et al. 2018) and with a latitudinal profile mostly distributed in the northern hemisphere between 10° and 45° of latitude (Bourgoin 2020). The family is divided into two easily distinguishable subfamilies: Flatinae Spinola, 1839 with fore wings, which are held parallel to the body, and Flatoidinae Melichar, 1901 with wings held horizontally (O'Brien 2002). In Madagascar, Flatidae are represented by 21 genera with 52 species of Flatinae and 11 genera with 41 species of Flatoidinae (32 genera and 93 species in total). Madagascan Selizini Distant, 1906 is represented by five genera and eight species. Species belonging to the genus Urana Melichar, 1902-U. paradoxa Melichar, 1902and U. unica Stroiński & Świerczewski, 2012 -are recorded across north-south mountain ranges and associated with the ecotone vegetation of high altitude-medium altitude moist evergreen forest (Stroiński & Świerczewski 2012 (Świerczewski & Stroiński 2019b). All genera listed above together with Stenocyarda angustata (Melichar, 1902) are endemic to the island.
In this paper, an additional monotypic genus of flatid planthoppers Selizitapia gen. nov., is described for Selizitapia pennyi gen. et sp. nov. (type species), which is associated with, endemic to Madagascar, a vegetation formation known as tapia woodlands.

Material
The material studied comes from the collection of the California Academy of Sciences in San Francisco, USA. For more than two decades, Academy entomologists have led research expeditions to the island and their results can be found under the following link: https://www.calacademy.org/learn-explore/scientific-expeditions/madagascar. Label information of all specimens examined is between double quotation marks and provided verbatim with each line separated by a slash (/).

Preparations and illustration
All specimens were originally preserved in 70% ethanol, but were subsequently dried and pinned. The abdomens of the examined specimens were removed and cleared for 30 minutes in warm (50°C) 10% KOH solution with a few drops of chlorazol black (CAS No. 1937-37-7) for staining the ectodermic structures based on the method introduced by Carayon (1969) and Bourgoin (1993). Dissections and cleaning of genital structures were performed in distilled water. Final observations and drawings of genital structures were done in glycerol using a camera lucida attached to a light microscope. The SEM photographs of dry, uncoated specimens were taken in the Laboratory of Scanning Microscopy, Museum and Institute of Zoology, Polish Academy of Sciences (Warsaw), using a scanning electron microscope HITACHI S-3400N under low vacuum conditions.
Descriptions of the external characters of head, thorax and abdomen were based on SEM photographs, whereas descriptions of male and female terminalia were based on macerated and stained structures.

Measurements and abbreviations
Measurements were made with an ocular micrometer. The following measurements, ratios and their abbreviations were used in this study: Total length = length of specimen from anterior margin of head to tegmina apex (in dorsal view) A/B = width of vertex at anterior margin / length of vertex at midline C/E = width of frons at upper margin / length of frons at midline D/E = maximum width of frons / length of frons at midline F/B = length of pronotum at midline / length of vertex at midline G/F = length of mesonotum at midline / length of pronotum at midline G/B+F = length of mesonotum at midline / cumulative length of vertex and pronotum at midline G/H = length of mesonotum at midline / width of mesonotum between lateral angles I/J = length of tegmen from the base to the apical margin in median portion / width of tegmen measured from the apex of clavus to the anterior margin I/K = length of tegmen from the base to the apical margin in median portion / width of tegmen at the widest part The metatibiotarsal formula LT-T/Mt1/Mt2 provides the number of spines on the side of the metatibia LT-on the apex of metatibia T, alternatively in two groups of internal Ti and external Te spines separated with a diastema (Ti-Te) / on the apex of first metatarsomere Mt1 / on the apex of second metatarsomere Mt2.

Terminology
The nomenclature of the forewing (tegmen) veins follows the interpretation proposed by Bourgoin et al. (2015). Antennal structures are named in accordance with Stroiński et al. (2011). The terminology of the genitalia follows Bourgoin (1988) and Bourgoin & Huang (1990) for the male, and Bourgoin (1993) for the female.

Etymology
The generic name is an arbitrary combination of the name of the tribe 'Selizini', which the new genus belongs to, and the name of the forest formation -'tapia', which the insect is associated with. Gender feminine.

Description
Head. Legs. Pro-and mesofemora slightly shorter than tibiae, subrectangular in cross section. Pro-and mesotibiae with shallow groove on external side; apical tarsomere of anterior and median legs longer than cumulative length of second and basal tarsomeres. Metatibiae longer than metafemora, triangular in cross section with two lateral spines and apical row of spines -first lateral spine placed subapically, second lateral spine placed a bit after midlength, apical spines in formula 2 longer (external) + 5 shorter (internal); basitarsomere of metatarsus a bit longer than cumulative length of second and apical tarsomeres, with apical spines lined as semicircle -2 external spines a bit longer than 7 shorter internal spines; each internal spine bearing single, distinct seta; second segment of tarsomere with two lateral spines and median pad with setae. Metatibiotarsal formula: 2-(2+5)/(2+7)/2 (Figs 3B-D).

Diversity and distribution
The genus is monotypic and contains a single species from Madagascar.

Diagnosis
The only species in the genus.

Etymology
The species is dedicated to Dr Norm Penny (

Paratypes
All specimens with the locality collection data the same as the holotype apart from the collection codes and the dates of collecting. All dry-mounted, abdomens of some specimens detached, dissected and stored in glycerol in a glass microvial under the specimen. coLoraTion. Mostly uniformly stramineous with costal area, apical cells and postclaval margin fuscous; dark brown markings between basal part MP and CuA; teeth of gonoplac dark brown (Fig. 1A-C). MaLe TerMinaLia. Anal tube, in lateral view, with ventral margin as obtuse angle and dorsal margin arcuate (Fig. 5A). Genital style with posterior margin weakly convex, ventral and dorsal margins almost straight, postero-ventral angle bluntly rounded, not extending to the posterior margin (Fig. 5A). Basal lobes of ventral periandrium small with sinuate margin (Fig. 5D). FeMaLe TerMinaLia. Pregenital sternite with anterior and posterior margins in median portion almost straight (Fig. 7A). Anal tube with apical margin, in dorsal view, strongly rounded (Fig. 7B); in lateral view, with basal part wider than apical part, anus placed almost at midlength (Fig. 7C). Gonoplacs with membranous part at ventral margin (Fig. 7D). Gonapophysis VIII with subapical 5 massive teeth at dorsal margin (Fig. 7E). Ductus receptaculi and diverticulum ductus widened distally (Fig. 7G).

Distribution and habitat
Madagascar; so far only known from one locality in the central part of the island (35 km SSE of Antsirabe). The type series was collected in a tapia woodland formation, in the season of 2003 from March to May and from September to October.

Discussion
The biodiversity of Madagascar can be characterised by a high rate of endemism in species and at higher taxonomical levels (Waeber et al. 2019). This may result from a broad ecological diversity occurring in limited areas, with a wide spectrum of unique niches, such as here the tapia woodlands, a humanmodified vegetation formation found in highland zones of central and southwestern Madagascar. It is characterized by two strata: a woody stratum dominated by Uapaca bojeri Baill. (tapia) in association with other woody species of the family Sarcolaenaceae (Madagascar's largest endemic family) and a multi-layer herbaceous stratum represented primarily by species of Asteraceae, Cyperaceae, Lamiaceae, Poaceae, and Rubiaceae (Alvarado et al. 2014a Protection of tapia woodlands equals protection of unique invertebrate communities associated with this formation. The existence of this anthropized vegetation results from the interaction of localised edaphic conditions, human activities and fire. The latter seems to be the most important factor, as was noticed by Alvarado et al. (2014b). The greater frequency and extent of intentionally set bush fires has led to greater diversity among woody and herbaceous native species. Selective exploitation also plays an important role in the conservation of tapia woodland. The economic importance of Uapaca bojeri fruits and of the woody species for wild silk worms should be factored into management planning. However, the most important threat to tapia woodlands is the invasion by exotic plants (Kull 2014).
The high endemism of Madagascan Selizini can be compared to that of Socotra Island where all genera of Selizini described so far seem to be known exclusively from this area. There, we can also observe a unique altitudinal distribution of particular species firmly associated with the different types of vegetation diversified according to the specific terrain relief and changing climatic conditions (Stroiński et al. 2016Świerczewski et al. 2017. However, similarly to Madagascar, all these biotopes are under permanent human-related threats.
In summary, taking into account the enormous loss of the forest cover in Madagascar and the degradation and fragmentation of its habitats, conservation management of these so particular biotopes should be undertaken to allow their protection at all scales, from landscapes to communities and populations. These are crucial to the future survival of herbivorous insect communities composed of highly-specialized species associated with particular native plants and influenced by specific abiotic factors.